Robert Galambos

8/4/2019 Robert Galambos

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EDITORIAL ADVISORY COM MITTEEAlbert J . AguayoBernice Grafstein

Theodore MelnechukDale Purves

Gordon M. Shep herdLa rry W. Swa nson (Chairperson)


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T h e H is to r y o f N e u r o s c i en c ein A uto b io g ra phyVOLUME 1

E dited by L arry R. Squire

SOCIETY FOR NEUROSCIENCEWashington, D.C.

1996


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Society for Neuroscience1121 14th Street, NW., Suite 1010

Washington, D.C. 20005

1996 by the Society for Neuroscience.

All rights reserved.

Printed in the United States of America.

Library of Congress Catalog Card Number 96-70950ISBN 0-916110-51-6


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Contents

Denise Albe-Fessard 2

Julius Axelrod 50

Peter O. Bishop 80

Theodore H. Bullock 110

Irving T. Diamond 158

Robert Galambos 178

Viktor Hamburger 222

Sir Alan L. Hodgkin 252

David H. Hubel 294

Herbert H. Jasper 318

Sir Bernard Katz 348

Seymour S. Kety 382

Benjamin Libet 414

Louis Sokoloff 454

James M. Sprague 498

Curt von Euler 528

John Z. Young 554


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R o b e r t G a l a m b o sBORN: Lorain , OhioApril 20, 1914EDUCATION: Oberlin College, B.A., 1935

H ar va rd Un iversi ty, M.A., Ph.D . (Biology, 1941)U niv ers i ty of Roche ster , M.D., 1946APPOINTMENTS"Harvard Medical School (1942)Em ory Un iver s i ty (1946)Harvard Univer s i ty (1947)W al t e r Reed Arm y Ins t i t u t e o f Research (1951)Yale Universi ty (1962)

U nive rs i ty of Cal i forn ia , Sa n Diego (1968)Pro fesso r o f Neurosc iences Em er i tus , Un ive r s i ty o fCal i fornia, San Diego (1981)HONORS AND AWARDS:Am erican Aca dem y of Ar ts and Sciences (1958)N at ion al A cadem y of Sciences U SA (1960)

Robe r t Galambo s d iscovered , w i th Do na ld Gr i ff in , t he ph eno me no n o fecho loca t ion i n ba t s . Dur ing h i s career he carr i ed ou t f undamenta l

phys io log i ca l s t ud i es o f t he aud i tory sy s t em us ing microe lec t rodes i n ca ts ,a n d l a te r s t u d i e d b r a i n w a v e s a n d a u d i t o r y e v o k ed p o t e n t i a l s i n h u m a n s .

He was an ear l y and force fu l pro ta gon i s t f or t he impor tance o f g l i ain t he f unc t ion o f the nervou s sys t em .


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R o b e r t G a l a m b o s

I n t r o du c t i o nT h e s u b j e c t w a s b o r n i n L o r a i n , O h i o , o n A p r i l 2 0, 1 9 1 4 , n o tl o n g a f te r t h e v a c u u m t u b e w a s i n v e n t e d . A t th e a g e o f 6 , a n di n t h e f i r s t g r a d e o f a C l e v e l a n d , O h i o p u b l i c s ch o o l, h e h e a r dh i s f i r s t r a d i o m e s s a g e t h r o u g h a n e a r p h o n e c o n n e c t e d t o ac r y s t a l r a d i o r e c e i v e r h i s o l d e r b r o t h e r h a d b u i l t . H e w a sa b o u t 4 0 y e a r s o l d w h e n t e l e v i s i o n s e t s f i r s t a p p e a r e d f o r s a l ei n t h e s t o r es ; b y t h a t t i m e h e h a d o b t a i n e d A .B . a n d M . A .de gr e e s i n Zoo l ogy a t Obe r l i n Co l l e ge ( 1936) ; M.A. a nd Ph .D .d e g r e e s i n B i olo g y a t H a r v a r d U n i v e r s i t y ( 19 4 1); a n d t h e M .D .d e g r e e a t R o c h e s t e r U n i v e r s i t y ( 1 9 4 5 ) . A l s o , p e n i c i l l i n h a db e e n d i s c o v e r e d , H i t l e r a n d H i r o h i t o d e f e a t e d , a n d a r e m a r k -a b le e x p a n s i o n o f r e s e a r c h o n th e b r a i n w a s j u s t g e t t i n g u n d e rw a y t h r o u g h o u t t h e w o r l d . T h i s e s s a y p r o v i d e s s o m e d e t a i l sa b o u t t h e s u b j e c t ' s p a r t i c i p a t i o n i n t h a t e f f o r t .

I n a u t o b i o g r a p h i e s t h i s u s e o f t h e t h i r d p e r s o n p a s t t e n s e i s t h e w a yw r i t e r s i n f o r m r e a d e r s t h e y f e e l u n c o m f o r t a b l e w i t h t h e t o p i c b e i n gd i s cu s s e d . M y p r o b l e m i s t h a t I h a v e a l r e a d y p u b l i s h e d o n e o f t h e s es e l f - p o r t r a i t s ( G a l a m b o s , 1 9 9 2 ) , w h i c h i s p r o b a b l y a l l t h e w o r l d n e e d s .H o w w i ll I c o v er t h e s a m e o l d g r o u n d i n a n e w w a y ? T h e q u e s t i o n s I a s k e di n s e a r c h o f t h e a n s w e r m a y b e w o r t h p r e s e rv i n g .

W h o w r i te s a n a u t o b io g r a p h y ? A m o n g m o d e r n s c ie n t is ts , a l m o s ti n v a r ia b l y , s o m e o n e w h o h a s b e e n a s k e d . B e n j a m i n F r a n k l i n , o u r f i rs tg r e a t s c i en t i st , w r o t e a l on g o ne , a n d A b r a h a m L i n c o ln w r o t e a v e r y s h o r to n e, b u t w e d o n ' t r e m e m b e r e i t h e r m a n b e c a u se o f w h a t h e w r o te a b o u th i m s e l f . I f w h a t y o u p r o d u c e d u r i n g y o u r l i fe t im e i s re a l l y w o r t h w h i l e o t h -e r s s e e t o i t t he wor l d doe s no t f o r ge t .

W h y d o e s a p e r s o n a g r e e t o w r i t e o n e ? If y o u h a v e g r a n d c h i l d r e n ,w h i c h m o s t a u t o b i o g r a p h e r s d o , t h e i m m o r t a l i t y y o u r g e n e s c l a m o r f o r i sa l r e a d y a s s u r e d . D u t y ? V a n i ty ?F o r w h o m d o w e w r i t e ? I h a v e y e t t o f i n d s o m e o n e w h o m a k e s t h i se x p l i c i t , b u t I w i l l a i m m y a u t o b i o g r a p h y a t t h e y o u n g p e r s o n a b o u t t os u b m i t a m a n u s c r i p t r e p o r t i n g h i s o r h e r f i r s t s u c c e s s f u l e x p e r i m e n t ,


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R o b e r t G a l a m b o s 1 81

k n o w i n g f ul l w e l l t h a t w h e n I w a s a t t h a t p o i n t in m y o w n c a r e e r t h e b o t-t o m m o s t i t e m o n m y r e a d i n g l i s t w a s a n a c c o u n t o f s o m e o n e ' s l ife .

W h a t s h o u l d I w r i t e a b o u t ? I a s k e d s e v e r a l f r ie n d s , a n d t h e i r a n s w e r sc l u s te r e d a r o u n d t w o t h e m e s . M a n y w a n t e d to k n o w h o w I d e c i d ed w h a tI w a s g o i n g t o d o, b o t h a s a s t u d e n t b e fo r e c o m m i t t i n g m y s e l f t o a r e s e a r c hc a re e r, a n d t h e n e v e r y m o r n i n g a s I o p e n e d t h e d o o r o f m y p l a c e o f b u s i -n e s s a n d w a l k e d i n s id e . S t u d e n t s o f te n r a i s e d p r a c t ic a l m a t t e r s , s u c h a sh o w t o w r i t e a g o o d s c ie n ti fi c p a p er , h o w m a n y m i s t a k e s a r e y o u a l lo w e dt o m a k e d u r i n g a c a r e e r, a n d s o o n.

I f i n al ly s e t t l e d o n w h a t f o ll ow s , w h i c h h a s t h r e e p a r t s : m y b a c k -g r o u n d , m y w o r k , a n d w h a t I w o u l d d o i n t h e f u t u r e i f I h a d o n e. I t is as t o r y a b o u t p e o p l e , i d e a s , w h a t w e a c c o m p l i s h e d t o g e t h e r , a n d t h e e n v i -r o n m e n t s i n w h i c h w e w o r k e d d u r i n g th e m o s t r e m a r k a b l e 6 0 y e a r s i n t h eh i s t o r y o f s c i e n c e , s o f ar .P e r s o n a l M a t t e r sI w a s t h e t h i r d o f f o u r b r o t h e r s . M y f a t h e r ( 1 8 80 - 1 95 4 ) a n d m o t h e r ( 18 8 5-1 96 9) c a m e t h r o u g h E l li s I s l a n d f ro m n o r t h e a s t H u n g a r y a r o u n d 1 89 5a n d m e t f o r t h e f i r s t t i m e i n L o r a i n a b o u t 1 0 y e a r s l a te r . M y p a t e r n a lg r a n d f a t h e r ( 1 84 4 -1 9 07 ) w a s a p e a s a n t w h o d ie d i n t h e s a m e f a r m h o u s ew h e r e h e h a d r a i s e d t w o d a u g h t e r s a n d f o u r so n s, o f w h o m m y f a t h e r w a st h e y o u n g e s t . ( I h a v e a c o py o f t h e v o n G a l a m b o s c o a t of a r m s a n d o n cee x c h a n g e d l e t t e r s w i t h t h e l a s t n o b l e m a n o f t h e l i ne ; t h e r e is n o e v id e n c ew h a t e v e r o u r f am i l i e s a r e r e la t e d . ) M y m o t h e r , J u l i a P e t i ( P e tt y ), w a s t h eo l d e s t of f iv e s ib l in g s ; h e r f a t h e r w a s a s c h o o l t e a c h e r w h o t a u g h t h e r t or e a d a n d w r i t e b e fo r e s h e w a s b r o u g h t to A m e r i c a b y a r e l a t i v e a t t h e a g eo f 1 2 o r 13. It is i n t e r e s t i n g a n d s a d t h a t I r e t a i n n o t h i n g t h a t I m a y h a v eb e e n to ld a b o u t m y g r a n d m o t h e r s .M y f a t h e r s a i d h i s f i rs t p u r c h a s e w a s a n E n g l i s h d i c t io n a r y , a n d t h a th e s e t h i m s e l f t h e t a s k o f l e a r n i n g t o s pe ll , p r o n o u n c e , a n d u s e t h r e e n e ww o r d s e v e r y d a y . B y 1 9 0 5 h e h a d a p p r e n t i c e d a s a c a r p e n t e r a n d w a s t a k -i n g a c o r r e s p o n d e n c e s c ho o l c o u r s e c o v e r in g t h e b u i l d i n g t r a d e s , a n dw o u l d s o o n s e t h i m s e l f u p i n t h e h o u s e - b u i l d i n g b u s i n e s s h e s u c c e s s f u l l yc o n d u c te d t h r o u g h o u t h i s life . H e w a s p r o u d t h a t h i s w o r d a n d h a n d s h a k ew e r e a l l a n y o n e n e e d e d t o c l o s e a b u s i n e s s d e a l .

M y m o t h e r w a s a s m a l l w o m a n - - p e r h a p s f iv e f ee t t a l l - - w h o t oo k n o n-s e n s e f r o m n o on e . S h e a t t e n d e d n i g h t s c ho o l t o i m p r o v e h e r E n g l i s hs k i ll s , a n d I r e t a i n d o z e n s o f l e t t e r s s h e w r o t e i n a c u r i o u s l y a n t i q u e h a n d .S h e t a u g h t h e r s o n s p r o m p t n e s s b e c a u s e t h e e a r ly b ir d c a tc h e s t h e w o r m ,f r u g a l i t y b e c a u s e a p e n n y s a v e d i s a p e n n y e a r n e d , a n d h o n e s t y b e c a u s ei t is th e b e s t p o l i c y - - t e a c h i n g s m a n y y o u n g p e o p le t o d a y n e v e r e v e ne n c o u n t e r , l e t a lo n e l e a r n . A s a n a d u l t I s p e n t a d a y o r t w o w i t h h e r w h e n -e v e r p o ss ib l e, m a n a g i n g t h i s o n c e o r t w i ce e v e r y y e a r a t h e r h o m e i n


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182 Robert GalambosF l o r id a . S h e t o ld m e d u r i n g m y l a s t v i s it s h e h a d d e l a y e d g o i n g t o t h e h o s -p i t a l b e c a u s e s h e w a s c e r t a i n s h e w o u l d c o m e o u t " f e e t f i r s t " t h e n e x tt im e . A n i n o p e r a b l e g a s t r ic c a r c i n o m a h a d f i n a ll y b l o ck e d h e r d i g e st iv et r a c t . M y m o t h e r w a s m u c h l o v ed ; t h r e e o f h e r d o c to r s h e l p e d c a r r y h e rc a s k e t a n d , a s sh e h a d i n s t ru c t e d , w e d r a n k c h a m p a g n e d u r i n g th e g o od -b y e p a r t y a t m y b r o t h e r ' s h o u s e a f t e r w a r d s .

M y p a r e n t s w e r e i n t e l l i g e n t b u t n o t i n t e l l e c t u a l s ; t h e r e w e r e f e w i fa n y f a m i l y d i s c u s s i o n s a b o u t b o o k s , r e l ig i o n , p o e t ry , o r p o li ti cs . M y f a t h e rd i d o n c e o u t l i n e f or m e h i s t h e o r y o f v i si o n ; i t i n v o l v e d p a r t i c l e s e m i t t e db y th e e y e t h a t r e a c h t a r g e t s i n th e e n v i r o n m e n t . M y m o t h e r l i s te n e dp r o u d l y w h i l e I d e s c ri b e d m y r e s e a r c h r e s u l t s , b u t s h e s ti ll w o n d e r e d h o ws o o n I w a s g o i n g t o g o t o w o r k w h e n I w a s a l m o s t 4 0 y e a r s o l d .Phy sical Well-Being; Fina ncial Security, Dom estic T ranq uilityP r i o r t o a m i l d h e a r t a t t a c k a t a g e 7 8 , m y m o s t s e r i o u s m e d i c a l p r o b l e m sh a d b e e n a t o n s i l l e c t o m y a t 1 9 , a f r e q u e n t l y a c h i n g b a c k , a n d a n o c c a -s i o n a l l y p a i n f u l k n e e c o r r e c t e d b y a r th r o s c o p i c s u r g e r y a t a g e 6 9 . A t 6 5 Iq u i t s m o k i n g a f t e r 5 0 y e a r s , b e g a n jo g g i ng , a n d k e p t a n a l m o s t d a i ly lo go f d i s t a n c e r u n f or t h e n e x t 1 0 y e a r s . I t s e n t r i e s o c c a s io n a ll y n o t e w h a t ag o d s e n d t h i s e x e r c i s e w a s f o r m e p h y s i c a l l y a n d m e n t a l l y , a n d t h e y a l s ot r a c e , i n a d v e r t e n t l y , t h e o r d e r i n w h i c h m y g e n e s h a v e p r o g r e s s i v e l yt u r n e d o f f o n e b o d i ly p r o c e s s a f t e r a n o t h e r . A t 8 1 I h a v e f i n a l l y a c c e p t e dt h e f a c t t h a t a fe w y e a r s a t m o s t r e m a i n f or c o m p l e t in g w h a t I s ti ll w a n tt o d o , a n d a m m i l d l y a m u s e d a t h o w , l i k e s o m a n y o t h e r a g i n g p e o p l e , Is t u b b o r n l y r e f u s e d to a c c e p t m y m o r t al it y .M o n e y h a s n e v e r b e e n m u c h o f a p r o b le m , a l t h o u g h I w a s c lo se t o 4 0b e f o r e r e p a y i n g w h a t I h a d b o r r o w e d f r o m p a r e n t s a n d o t h e r s .T h r o u g h o u t m y a d u l th o o d , t h e n a t i o n a l e c o n o m y e x p a n d e d , s a la r ie si n c r e a s e d r e g u l a r ly , a n d i n f l a t i o n b o o s t e d t h e v a l u e o f t h e h o m e s I so ld . A sa r e s u l t , I f o u n d i t p o s s i b le t o li v e w e l l w i t h m y f a m i l y a n d t o d o s u c h e x t r at h i n g s a s p a y t h e s a l a r y o f a c o ll a b o r at o r f or a m o n t h o r tw o b e t w e e ng r a n t s , c o m m u t e to B u d a p e s t to w o r k w i t h c o l le a gu e s o n a n e x p e r i m e n t ,a n d a s s e m b l e a co l le c ti o n o f o ld p o c k e t w a t c h e s a n d N a v a j o r u g s .

I h a v e h a d t h r e e w i v e s , e a c h a s t r o n g p e r s o n w h o m e s h e d h e r c a r e e rp l a n w i t h m y o w n . M y f i r s t w i f e , n o w J e a n n e t t e W r i g h t S t o n e , i s w i d e l yk n o w n f o r h e r c o n t r i b u t i o n s t o t h e f i e ld o f e a r l y c h i l d h o o d e d u c a t i o n ; a f t e rm o r e t h a n 3 0 y e a r s , s h e c h o se to d iv o rc e m e fo r a n o t h e r m a n . T h e s ec o n d ,C a r o l A r m s t r o n g S c h u l m a n , a n e u r o s c i e n t i s t i n h e r o w n r i g h t , l ef t m e b yc o m m i t t i n g s u i c id e d u r i n g o n e o f h e r b o u t s o f d e p r e s si o n . T h e t h i r d ,P h y l l i s J o h n s o n , j o i n e d m e i n 1 9 7 7 , a n d s i n c e t h e n I h a v e k n o w n m o r ep e a c e , o r d e r , c o m f o r t , a n d c o m p a n i o n s h i p t h a n a p e r s o n h a s a n y r i g h t t oe x pe ct . J e a n n e t t e a n d I h a v e t h r e e d a u g h t e r s , w h o , b e t w e e n t h e m , h a v eg i v e n u s f iv e g r a n d c h i l d r e n .


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Ro b e r t G a l a m b o s 183A w a r d s a n d P r i z e sM y h o n o r a r y d e g r e e s i n c l u d e t h e M .A . ro u t i n e l y a w a r d e d a l l Y a le p r o fe s -s o rs w h o d o n o t a l r e a d y h a v e a Y a le d e g r ee a n d t h e M . D . a w a r d e d b y th eS w e d i s h U n i v e r s i t y o f G S t e b o rg , fo r w h i c h i ts t h e n - r e c t o r , m y f r i e n dH o l g e r H y d e n , t h e g l i a sp e c i a li s t, i s p r o b a b l y r e s p o n s i b l e . I al s o h a v e s ev -e r a l m e r i t o r i o u s p e r f o r m a n c e c o m m e n d a t i o n s f ro m t h e A r m y . A f o r m e rc o ll ea g u e, G e o r g e M o u s h e g i a n , t o ld m e r e c e n t l y t h a t d u r i n g t h e p a s t 2 0 to3 0 y e a r s h e h a s r e p e a t e d l y s u b m i t t e d m y n a m e f o r v a r i o u s h o n o r s g i v e ns p e c ia l is t s i n h e a r i n g m a t t e r s a n d i s f r u s t r a t e d t h a t n o n e h a s e v e r b e e na w a r d e d . P e r h a p s h e o v e r e s t i m a t e s m y q u a l if ic a t io n s , b u t c e r t a i n ly m ya b i l it y to s a y n o w h e n o ff er ed j o bs t h a t w o u l d t a k e m e a w a y f r o m t h e l ab -o r a t o r y h a s p l a y e d a p a r t . M y o w n v i e w i s t h a t I a m o f t e n a r r o g a n t a n dc r a n k y , a n d t h i s t u r n s p e o p l e o f f .In t ro du ct ion to Research , Ob er l in College , 193 4-38I f i r s t sys temat ica l ly encountered b io log ica l fac t s and concepts as a co l legejun io r i n 1934 and found them su rp r i s i ng ly ea sy t o g r a sp , r emember , andm a n i p u l a t e . M y m a t h a n d p h y si c s g r a d e s w e r e B w i t h a s p r i n k l i n g of C . I w a sde l igh ted by my spec ia l knack for Bio logy , which in re t rospec t seems easy toin t e r p re t i n t he con t ex t o f Ho wa rd G ardn e r ' s i dea o f m u l t i p l e i nna t e i n te l li -gences (Ga rdne r , 1983) . U nd e rg rad ua t e s i n 1935 were s t rong ly i nc li nedt o w a r d J .B . W a t s o n 's b e h a v i o ri s m , s o m e t i m e s i l lu s t r a t e d b y t h e f a n t a s y t h a ta g i v en b a b y c a n b e f as h i o ne d i n to e i t h e r a m u s i c i a n o r a m a t h e m a t i c i a n b yse lec t ing the prop er s t im ul i to c rea te i t s repe r to i re o f re f lex responses .

T h e c o n c e p tu a l d i s ta n c e i s i m m e n s e b e t w e e n s u c h i d ea s a n d t h e c u r r e n texp lana t i ons , wh ich a s s ign a huge con t r i bu t ion t o t he genom e ( "na tu re" ) andw h a t e v e r r e m a i n s t o " n u r tu r e . " G a r d n e r ' s S e v e n I n te l li g e n ce s a c c o u n t m u c hmo re ap t l y t ha n J .B . W a t son 's r e fl exes for t he mu s i ca l gen iu s o f M oza r t an dB a c h, t h e m a t h e m a t i c a l g e n i u s o f T u r i n g a n d L e i bn i tz , t h e v e r b a l g e n i u s o fShak espea re , an d t he a th l e t i c gen iu s o f ba l l e t dance r s an d bask e tba l l p l ay -e r s. I t s eem s be l ievab l e t o me tha t e ach o f u s a r r i ve s w i th a un ique mix o fG a r d n e r ' s s e v e n , a n d w e t h e r e a f t e r d e ve lo p th e s e to t h e e x t e n t p e r m i t t e d b ywh ere , a nd how long , we ha pp en to live. Of course , people s t i ll t ake s ides ont h e n a t u r e - n u r t u r e d ic h ot om y , b u t m y q u a i n t b e h a v i o r i st ic v ie w d i s a p p e a r e dforever fo l lowing the pub l ica t ion by J . D . Watso n a nd Cr ick , in 1953 , o f w ha tW atson ha s ca l led the i r " ins igh t in to the na tu re o f l ife i tse lf ."About t he Sc i en t i f i c PaperMy f i r s t encoun te r w i th one o f t he se t ook p lace i n my jun io r ye a r i n t hed e p a r t m e n t a l l i b r a r y a s I w a s p r e p a r i n g m y f i r s t s e m i n a r r e p o r t f o r C . G .Rogers , a p rofessor o f C om pa ra t iv e Phys io logy . The paper , by W. R. Hess ,


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184 Rober t Ga lambosd e a l t w i t h t h e n e r v o u s s y s t e m o f t h e e a r t h w o r m , a n d i t e n d e d w i t h a c o m -p l e te s u m m a r y o f t h e p a p e r ' s o b je c ti ve s a n d r e s u l ts ! W h a t a s t u n n i n g s u r -p r is e ! How in fo rm a t ive and he lp fu l! I gushed on li ke th i s i n m y p re sen t a t i on .I have neve r r ea d an accoun t o f how the s c ien ti fi c pape r , t ha t un iqu e c re -a t ion of the sc ien t i fi c com mu ni ty , evo lved to reach i t s mo de rn form. Thema thema t i c i an Mark Kac once ca l l ed t hem f ive -pa r t Sc i en t i f i c Sona t a s :Sum m ary , I n t roduc t ion , M e thods , Resu l t s , D i scus s ion . I t i s c lea r ly t he be s tkno wn w ay to o rgan ize a s c i en t if ic message ; t r y t o i nven t s om e th ing d i f f e r en tand be conv inced . Meanw hi l e , he r e a r e two t i p s i f you need he lp : f i rs t , s t ud ya f ew pub l i shed example s you admi re and no t e how o f t en t he wr i t e r s f o l l owthe ru l e s you w i l l f i nd i n S t runk and Whi t e ' s The Elements o f S ty le . Second ,e d i t r u th l e s s ly ; y o u c a n a l w a y s i m p r o v e w h a t y o u h a v e a l r e a d y w r i t t e n .M y Fi rs t Labora toryR a y m o n d H e r b e r t S t e t s o n , p r o f e s s o r o f p s y c h o lo g y a t O b e r l i n C o l le g e i n1 9 35 , w a s o n e o f t h o s e u n s u n g h e r o s o f A m e r i c a n s ci en c e: t h e s m a l l -c o l le g ep r o f e s s o r w h o i n s p i r e s a n d g u i d e s i t s r e c r u i t s a t t h e t i m e t h e y a r e m o s tv u l n e r a b l e a n d e d u c a b l e . H e i n t r o d u c e d m e t o t h e r e s e a r c h p l a n , t h er e s e a r c h l ab , a n d t h e r e s e a r c h d is co v er y. I n m y tw o y e a r s w i t h h i m( S e p t e m b e r 1 93 5 -3 7) I l e a r n e d a ll t h e f u n d a m e n t a l s : h o w to f o r m u l a t e t h ep r o b l e m , p l a n t h e w o r k , c o ll ec t t h e a p p a r a t u s , d o t h e e x p e r i m e n t s , a n a l y z et h e d a t a , m a k e t h e f i g u r e s , w r i t e t h e p a p e r , g e t i t p u b l i s h e d , a n d , f i n a l l y ,h o w t o t e a c h w h a t y o u k n o w a b o u t a l l t h i s t o o t h e r s . S e e K e l s o a n dM u n h a l l ( 19 88 ) fo r b i o g r a p h ie s o f t h i s r e m a r k a b l e m a n .R o g e r S p e r r y a n d I g r a d u a t e d t o g e t h e r i n 1 9 35 a n d t h e n d id o u r m a s t e r ' sdeg ree r e sea rch i n S t e t son ' s Osc i l l og raph Labora to ry , wh ich , t hanks t o i t sch i e f t e chn ic i an , J am es M. Snodgra s s , w as ab ou t a s w e l l equ ipped for el ec -t ro p h y si o lo g i ca l m e a s u r e m e n t s a s t h e F o r b e s- D a v is H a r v a r d M e d i c a l Sc ho oll abo ra to ry t o wh ich I wou ld sho r t l y go. S t e tson ' s l ab r egu l a r ly i nc luded a f ewsen io r v i s it o r s who had come to work on t he m ech an i s m s o f speech p roduc -t ion , o r m o to r phone t i c s , S t e t son ' s spec ia l fi eld o f i n t e r e s t . I t was t he r e t h a t Ij o ined t he f i r st o f m an y such sma l l , i n t im a te f e l lowsh ips t h a t un i t e for t hepu rpose o f d is covery. Me m ber s o f eve ry h ea l t h y l ab bo nd c lo sely t oge the r , l i kea l l comrades who seek the same goa l . Years l a te r , a t Yale , I c rea ted s imi la rt em po ra ry g roups by o rgan i z ing sum mer - long , s i x -days -a -week oppo r tun i t i e s( f i ve i n a l l ) whe re young peop le ga ined hands -on expe r i ence w i th e l ec t ro -phys io log i ca l i n s t rumen t s and deve loped a ce r t a in sk i l l i n u s ing t hem. S t i l ll a t e r, i n Sa n D iego , t h i s becam e a t h r ee - t o fou r -day an nu a l sym pos i a ( s eveni n a ll ) o n th e t h e n - n e w a u d i t o r y b r a i n s t e m r e sp o n s e; t h e a t t e n d e e s l i st e n e dt o l ec t u re s , b u t m o r e i m p o r t a n t l y t h e y c a r r i ed h o m e t r a c i n g s o f t h e r e s p on s -e s m a d e w i t h t h e i r o w n h a n d s . I h a v e a l w a y s w a n t e d m y o w n l a b o r a t o r y t obe l i ke S t e t son ' s , a p l ace whe re peop le t ake p l ea su re i n c r ea t i ng t he i r ownexp e r im en t s and d i scove ri e s i n t he co mp any o f o the r s do ing t he s am e .


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Robert Galambos 1 85D o n L i n d s l e y i n t r o d u c e d m e t o s in g l e u n i t s w h e n h e v i s i t e d S t e t s o n 's l a b

in th e sp r in g o f 1 9 36 . W h i l e a t H a rv a r d (1 93 3 -3 5 ), D o n h a d in se r t e d e l ec-t ro d es c o n s i s t i n g o f a fi n e in su la t ed w i re in s id e a h y p o d e rm ic n eed le in to a rma n d l e g m u s c l e s o f h u m a n s u b je c t s to i s o la t e s i ng l e m o t o r u n i t s , w h i c h h ed e f in ed a s t h e co l l ect io n o f mu sc le f ib r e s i n n e rv a te d b y a s in g le mo to r n e u ro n .S t e t s o n h a d h e a r d o f t h e s e m e a s u r e m e n t s a n d a s k e d L i n ds le y , w h o b y t h e nw as a t t h e Wes te rn R ese rv e Med ica l S ch o o l i n C lev e lan d 3 0 mi l e s aw ay , t oc o m e a n d d e m o n s t r a t e h i s t e c h n i q u e . D o n a r r i v e d a n d c o n n e c t e d h i s e l e c -t ro d es to S n o d g ra s s am p l i f i e rs , w h i l e t h e l ab g ro u p (S p e r ry , Jo e M i ll er , H . D .B o u man , an d I ) w a tch ed . I c an s t i l l h ea r t h o se in d iv id u a l l o u d p o p s th e lo u d -s p e a k e r e m i t t e d , w h i c h D o n p r e d i c t a b l y a d j u s t e d d o w n a n d u p i n r a t e b yex e r t in g l e s s o r mo re e f fo r t. I n S te t so n ' s o p in o n , " mo to r u n i t " m ea n t o n e o f t h eo p p o s in g m u s c l e g r o u p s r e c i p r o ca l ly a c t i v a t e d a r o u n d s o m e j o i n t t o p r o d u c ea b a l l i s t i c mo v emen t , an d L in d s l ey ' s d i f f e r en t d e f in i t i o n t ro u b led h im. B u tS p e rr y , w h o s e m a s t e r ' s t h e s i s e x p e r i m e n t s m a p p e d t h e s e q u e n c e o f t h e s h o u l -d e r g i rd l e m u s c l e a c t i v a t i o n s d u r i n g s u c h b a l li s ti c m o v e m e n t s , w e l c o m e d t h en e w t e c h n i q u e s a n d i d e a s L i n d s l e y b r o u g h t .

A f t e r L i n d s l e y ' s v i s i t , S p e r r y a n d I f a b r i c a t e d c o n c e n t r i c n e e d l e e l e c -t r o d e s a n d i n v e n t e d n e w o n e s , t h e m o s t s u c c e s s f u l o f w h i c h w a s a s t r a n do f f i n e c o p p e r w i r e w i t h a s i n g l e l i n e c u t a c r o s s i t s i n s u l a t i o n w i t h as c a l p el b l a d e. W e t h r e a d e d t h i s w i r e i n to t h e e y e o f a s u r g i c a l n e e d l e ,p a s s e d t h e n e e d l e t h r o u g h o u r s k i n i n t o a m u s c l e a n d b a c k o u t , a n d c o n -n e c t e d i t t o t h e S n o d g r a s s a m p l i f i e r a n d l o u d s p e a k e r . W h e n o u r m u s c l ec o n t r a c t i o n s c a u s e d t h e l o u d s p e a k e r t o e m i t l o u d p o p s , s i m i l a r t oL i n d s l e y ' s , w e k n e w t h e b a r e d s u r f a c e r e s t e d u p o n o n e o r a f e w m u s c l ef i b e r s . I a l s o f o u n d t h a t a n o r d i n a r y b r a i n w a v e e l e c t r o d e p l a c e d o n t h es k i n o v er t h e f i r st d o r s a l i n t e r o s s e o u s m u s c l e - - t h e o n e c o n n e c t in g t h u m ba n d f o r e f i n g e r - - w i l l r e a d i l y p i c k u p s i n g le u n i t s i f o n e c a r e f u l ly a d j u s t st h e t e n s i o n e x e r t e d .M y m a s t e r ' s t h e s i s p r o p o s a l to th e z oo lo gy d e p a r t m e n t w a s t h e a n a l y -s is o f e a r t h w o r m l o co m o t io n u s i n g m u s c l e a c t io n c u r r e n t s r e c o r d e d i nS t e t s o n ' s l a b . S t e p o n e w a s t o b u i l d a d i r e c t c o u p l e d a m p l i f i e r ; S n o d g r a s sd e s i g n e d i t , I b u i l t i t , a n d i t s u c c e s s f u l l y a m p l i f i e d t h e p o t e n t i a l s a s s o c i -a t e d w i t h e a r t h w o r m m o v e m e n t s , w h i c h w e d i s p l a y e d w i t h b o t h aW e s t i n g h o u s e o s c i ll o g ra p h a n d a s m o k e d d r u m k y m o g r a p h . M y t h e s i s w a sa c c e p t e d i n 1 9 36 , b u t i t fe ll f a r s h o r t o f w h a t I h a d i n m i n d . S t e t s o n a g r e e dt o m y r e m a i n i n g a n o t h e r y e a r , a t t h e e n d o f w h i c h , s t il l d i ss a t is f i e d , Iw r o t e m y f i rs t p a p er , w h i c h w a s p u b l i s h e d i n th e F e s t s c h r i f t h o n o r i n g h i mo n h i s r e t i r e m e n t ( G a l a m b o s , 1 9 39 ).

T h r o u g h o u t m y s i x - y e a r O b e r l i n s t a y I p l a y e d s a x o p h o n e i n a d a n c eb a n d t o h e l p p a y m y b il ls , a n d w h e n I l e ft in t h e f al l of 1 9 37 f o r H a r v a r dw i t h t h e f e ll o w s h i p t h a t m a d e g o i n g t h e r e p o s s i bl e , I w a s a m e m b e r o f t h em u s i c i a n ' s u n i o n a b a n d o n i n g a p o s si b le m u s i c a l c a r e e r fo r w h a t I t h o u g h tw a s g o i n g t o b e t h e l i fe o f a s m o o t h - m u s c l e p h y s i o l o g i s t .


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186 Robert GalambosI n t r o d u c t i o n to N e u r o s c i e n c e , H a r v a r d I , 1 9 3 8 - 4 2At the Bio logica l Labora tor ies I met my advisor , A. C. Redf ie ld , a d is t in-gu i shed phys io log i s t and oceanographe r , and we wo rked ou t a cou r se o f s tudyth a t i nc luded m yco logy and i t s de l igh tfu l t eache r , C ap W es ton , and physio lo -g y, w h e r e G e o rg e W a l d m a d e m e m o r a b l e c o m m e n t s s u c h a s " t h e N a p o le o n o fsme l l has ye t t o be bo rn , " wh ich I gues s m ay s t il l be t rue . Redf i eld gave mean o ffice wh ere I s e t up s imp le in s t r um en t s fo r m eas u r in g the dynam ic p rop -e r t i e s o f i nve r t eb ra t e sm oo th musc le s , and l a t e r a r r an ged for me to spend thesu m m er o f 1939 a t t he B io log ica l S t a t ion in Be rm ud a where m y second, andla s t , con t r ibu t ion to smo o th m usc le phys io logy o r ig ina t ed (Ga lambos , 1941a ).

I t o l d R e d f i e l d v e r y e a r l y a b o u t m y i n t e r e s t i n e l e c t r o p h y s i o l o g y , a n dw i t h h i s b l e s s i n g v i s i t e d t h e F o r b e s - D a v i s H a r v a r d M e d i c a l S c h o o l l a b o -r a t o r y f or t h e f ir s t ti m e d u r i n g t h e 1 9 37 -3 8 w i n te r . A l e x a n d e r F o r b e s a n dH a l l o w e l l D a v is w e l c o m e d m e w a r m l y , a n d b e f o re l o n g I w a s m a k i n g t h et r i p f r o m C a m b r i d g e t o B o s t o n a t l e a s t o n c e a w e e k t o s e r v e a s a s u b j e c ti n E E G e x p e r i m e n t s , o r to w a t c h o t h e r e x p e r i m e n t s u n d e r w a y , a n d e v e nt o l e n d a h a n d f r om t i m e t o ti m e .

I n t h e l a t e 1 9 3 0s th e H a r v a r d M e d i ca l S ch o ol p h y s io l o g y d e p a r t m e n tw a s o n e o f a v e r y s m a l l n u m b e r o f p l a ce s i n t h e w o r ld w h e r e s t u d e n t s c o u ldl e a r n e l e ct ro p h y s io l o g ic a l t e c h n i q u e s . F o r s e v e r a l y e a r s F o r b e s a n d D a v i sh a d a g g r e s si v e ly s u p p o r t e d d e v e l o p m e n t o f t h e v a c u u m - t u b e a m p l if ie r s a n ds t i m u l a t o r s t h a t w e r e p r o p e l l i n g t h e d e p a r t m e n t i n t o t h e m o d e r n e r a o fb r a i n a n d p e r i p h e r a l n e r v e e le c tr o ph y s io l og y . A l b e r t G r a s s , w h o d e s i g n e da n d b u i l t a l l t h e p h y s io l o gi c al a m p l i f ie r s a n d s t i m u l a t o r s I u s e d , s u c c e e d e dE . Love t t Garceau , who had bu i l t t he l abo ra to ry ' s f i r s t ca thode r ay osc i l l o -g r a p h a n d E E G m a c h i n e . A l b e r t a r r i v e d a y e a r o r t w o be f or e I d id , a n d l e f ti n t h e e a r l y 1 9 4 0 s t o f o u n d h i s f a m o u s G r a s s I n s t r u m e n t C o m p a n y .

Seve ra l g r adu a te s tude n t s and pos tdoc to ra l f el lows were m eas u r in g b ra inwav es , evoked a nd cochlear poten t ia l s , an d s ingle ce ll respon ses ( I reca l l A. J .Derbysh i r e , J . E . H awk ins , J r. , H .O . Pa r r ac k , B . Renshaw, and P .O . The rman ) .B i rdsey R ensh aw showed m e m y f i r st f lu id - fi ll ed g l a s s p ipe t t e e l ec t rodes an dexp la ined how he h ad used th em to reco rd re sponses o f s ing le h ippocam pa lbra in ce l l s in situ (Renshaw et a l . , 1940); he lef t , his thesis f inished, short lyaf te r I a r r ived . His equipment passed f i r s t to a pos tdoctora l fe l low f romSweden , P . O . Th e rm an , t o whom Forbes app ren t i ced m e in the 1938-39 win -t e r. I i nhe r i t ed th i s s e t -up an d u sed i t w i th H a l Dav i s t o p roduce da t a fo r t hef i rs t tw o o f o u r t h r e e p a p e r s o n th e c o c hl ea r n u c l e u s - t h e o n e s e rr o n e o u s lyca l led audi tory nerve s tudies (Galambos and Davis , 1943; 1944) . Our th i rdpap e r i s a d i scl a imer , fou r yea r s l a te r , t ha t showed m an y o f ou r e l ec trodesm us t hav e been loca t ed in t he coch lea r nuc leus (Ga lamb os and Dav i s, 1948) .To the de t a i l ed accoun t o f t hese expe r imen t s wh ich appea r s e l s ew here(Galam bos , 1992a) , I would ad d o nly the fo llowing advice to the e age r grad u-a t e s tud en t o r pos tdoc a t an ea r ly s t age o f h i s o r he r c a ree r i n neu rosc i ence :


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R o b e r t G a l a m b o s 187D o e x a c t ly w h a t I d id . F i n d y o u r s e l f w e l c o m e d i n t o a l a b o r a t o r yw h e r e , f o r t h e f i rs t t i m e , o n e o f t h e m o s t i m p o r t a n t t e c h n i q u e so f t h e c e n t u r y h a s ju s t b e e n s h o w n t o w o rk . L e a r n to u s e t h em e t h o d f r o m i ts p i o n e er s . T h e n l i s te n c a r e f u l ly a s t h e l a b o r a to -r y d i r e c to r t e l ls y o u t h e s p a c e a n d e q u i p m e n t w i l l b e e x c lu s iv e -l y y o u r s i n t o t h e i n d e f i n i t e f u t u r e , a n d i n s t r u c t s y o u t o m a k ew h a t e v e r m e a s u r e m e n t s y o u w i sh . Y o u r s u c ce s s is a s s u r e d p ro -v i d e d y o u r e m a i n , o r b e c o m e , d il ig e n t a n d a t t e n t i v e .

Micropipet te E lec trodesI know of no scho la r ly h i s tory of the g la s s p ipe t t e mic roe lec t rode , b ut one ormo r e m a y i n f a c t ex i s t (S t e t son ga u ge d t he goodne ss o f a pa pe r by t he q ua l i t yof i t s l i t e ra tur e r eview) . Don Lind s ley says the F orbes -Davis l ab d id not havet he m whe n he l e f t t he p l a c e i n 1935 , bu t t wo ye a r s l a t e r i t c e r t a i n l y d i d ,be c a us e For be s , Re ns ha w, a nd Re mpe l de s c r i be d e xpe r i me n t s us i ng t he m a tthe 1938 me e t ing of the Am er ican P hys io logica l Soc ie ty (Re nsh aw e t a l ., 1938).

R e n s h a w ' s p i p e t t e s w e r e " p u l l e d b y h a n d o r w i t h a m a c h i n e d e v i s e da n d k i n d l y l o a n e d b y D r. L . G . L i v i n g s t o n f r o m t h o r o u g h l y c l e a n p y r e x c ap -i l l a r y t u b i n g . " A f t e r b r e a k i n g t h e 3 - 5 m i c r o n t i p s t o s i z e s " u p w a r d f r o m1 5t ," h e f i ll ed t h e m b y s u c t i o n w i t h a w a r m a g a r - s a l i n e s o l u ti o n , i n s e r t e da c h l o r i d e d s i l v e r w i r e i n t o t h e c o o l e d a n d h a r d e n e d a g a r , a n d d r o v e t h ee l e ct ro d e w i t h a m a n i p u l a t o r i n t o t h e b r a i n ( co r te x , h i p p o c a m p u s ) o f a n e s -t h e t i z e d o r d e c o r t i c a t e d r a b b i t s o r c a t s , a n d c h i c k e n e m b r y o s ( R e n s h a w e ta l . , 1 9 4 0 ) . H i s m i c r o e l e c t r o d e m e a s u r e m e n t s m a y b e t h e f i r s t e v e r m a d ei n s i d e a l iv i n g b r a i n . I n 1 9 3 9, u s i n g R e n s h a w ' s t e c h n i q u e , I p r e p a r e d i d e n -t ic a l p i p e t t e s w i t h 3 - 5 m i c r o n t ip s , f il le d t h e m b y s u c k i n g R i n g e r ' s s o l u t i o nu p i n t o t h e m u s i n g a 2 0 c c s y r i n g e w i t h i t s p l u n g e r c o a t e d w i t h V a s e l i n e ,a n d i n s e r t e d t h e m i n to t h e c o c h l e ar n u c l e u s a r e a o f a n e s t h e t i z e d c a ts .

R a l p h G e r a r d c l a i m s t o h a v e d i s c o v e r e d , i n 1 9 3 6 w i t h J u d i t h G r a h a m ,t he " t r ue mi c r oe l e c t r ode " whi c h he de f i ne s a s "a s a l t - f i l l e d c a p i l l a r y w i t h at i p s ma l l e nough ( up t o f i ve mi c r ons ) t ha t a mus c l e f i be r c ou l d be i mpa l e dw i t ho u t e xc e s s ive da m a ge " ( G e r a r d , 1975, p 468). The 1940 h i s t o r i c a l re v i e wi n Re n s h a w e t al . r e f e r e nc e s t he m i c r oe l ec t r ode s o f Ge l f a n da t e d 1927 , a ndof E t t i c k a nd Pe t e r f i da t e d 1925, a m on g o t he r s , bu t , c u r ious ly , no t t heG e r a r d a n d G r a h a m v e rs io n . A r e p r i n t T e d M e l n e c h u k r e c en t ly s e n t m edesc r ibes a 3-mic ron sa l ine - f i l l ed p ipe t t e used in 1918 by i t s au thor , I .H.Hyde ; s h e c a l ls he r s a mod i f i c a t ion o f one de s c r i be d i n 1910 by Ch a m be r s ,wh i c h in t u r n w a s ba s e d on t he e ve n e a r l i e r one o f M.A. Ba r be r ( Hyde , 1921).

G e r a r d , i n s u m m a r i z i n g h i s c a r e e r , s a y s " I a m p r o b a b l y b e s t k n o w nf o r t h e m i c r o e l e c t r o d e " ( G e r a r d , 1 9 7 5 p 4 7 4 ) . N o t b y m e . I r e m e m b e r h i mf or t h e r e m a r k a b l e G e r a r d , M a r s h a l l , a n d S a u l p a p er , t h e f i rs t c o m p r e -h e n s i v e e x p l o r a t io n o f t h e c o rt ic a l e v o k ed p o t e n t i a l s R i c h a r d C a t o n f ir s tde s c r i be d i n 1875 ( Ge r a r d e t a l . , 1936) .


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188 Robert GalambosBats

The collaboration with fellow gra dua te student Donald R. Griffin th at pro-duced my thesis experiments took place between May 1939 and November1940, sandwiched in between a summer in Bermuda and the Davis audi-tory microelectrode studies. It yielded six papers covered in my earlierautobiography (Galambos, 1941b;1942;1943a,b; Galambos and Griffin,1942; Griffin and Galambos, 1941). A paper just published adds details ofpossible historical interest (Galambos, 1995a) and I am happy to say werecent ly found the sound movies of flying bats tak en in 1941, thought forma ny years to be lost forever. Some advice: if your experiment is photo-genic, take the pictures and remember where you stored them afterward.By 1940, investigators had tried vainly for 150 years to discover themechanism by which blind bats avoid obstacles when flying. Today, in hind-sight, it is easy to identify the two completely unrelated technical advancesthat made the solution inevitable. One was the cochlear microphonicmethod for testing animal hearing, which Hal Davis was teaching me; theother was the development of the instruments t ha t generate, detect, andanalyze high-frequency sounds inaudible to man. Don Griffin, a graduatestudent already an authority on bats, had just published a paper reportingthey ut te r high-frequency cries inaudible to man; his co-author, G.W. Pierce,a physics professor, had just invented the ultrasonic sound generating andrecording instruments essential for the demonstration. Don asked me totest bat ears with the Davis method and within a month I had convincedmyself that the bat's upper hear ing limit was an octave or more above tha tof other animals. Don and I then designed and performed the behavioralexperiments that convinced us we had solved the problem. My recent his-torical account of those experiments concludes as follows:

Griffin and I were lucky, first of all, to have found each other,for it is not likely tha t either of us would or could have madethe measurements alone. Then there are the facts that the lab-oratories of Professors Pierce and Davis were separa ted by afew miles, and that their doors opened wide to us the momentwe knocked. And finally, every one of our experiments workedout exactly as planned, and they all pointed directly at the earhypothesis Jurine, and then Spallanzani, knew to be correct(in 1795 they both agreed that bats with plugged ears collidewith obstacles, but neither could say why this was so). At themoment we were united with our professors there was onlyone place in the world where two graduate students coulddemons trate tha t flying bats emit sounds we cannot hear, andthat the animals hear and act upon the echoes~and we hap-pened to be there (Galambos, 1996).


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R o b e r t G a l a m b o s 18 9A g r a d u a t e s t u d e n t o n ce a s k e d h o w I f o u nd t h e b a t p r o b l e m t h a t

b e c a m e m y P h . D . t h e s i s . F r a n k l y , I c a n n o t d e c i d e w h e t h e r I f o u n d t h ep r o b l e m o r t h e p r o b l e m f o u n d m e . I f a v o r t h e e x p l a n a t i o n t h a t c o u n t l e s si n t e r r e l a t e d e v e n t s , a c c u m u l a t e d o v e r 1 50 y e a r s , f i n al l y c o n v e r g e d o n t h et w o o f u s , a n d t h a t w e , li k e b u b b l e s i n th e v o r t e x t w i s t i n g a r o u n d t h e d r a i no f a n e m p t y i n g b a t h t u b , s w i rl e d f a s te r a n d f a s t e r a l o n g w i t h S p a l l a n z a n i ,o u r t w o H a r v a r d p r of e ss o rs , a n d t h e m a n y o t h e r s w h o h a v e le ft a m a r k i nt h e l i t e r a t u r e . I n t h i s f ig u r e o f s p e e ch , t h e p r o b l e m d i s a p p e a r s w h e n t h ed r a i n e m p t i e s ; h o w e v e r , 5 5 y e a r s l a t e r , a c c o r d i n g t o m y M e d l i n e s e a r c h ,a b o u t 2 5 b a t h e a r i n g p a p e r s a r e b e i n g p u b l i s h e d e v e r y y e a r .A l e x a n d e r F o r b e sI t i s n o t e a s y t o f i n d w o r d s t o d e s c r i b e t h e e n o r m o u s c h a n g e s i n r e s e a r c hm e t h o d s m y g e n e r a t i o n h a s s ee n . L e t m e t r y w i t h t h e s t o ry o f h o w o n e ofm y m e n t o r s , A l e x a n d e r F o r b e s , c a m e t o w o r k , a n d t h e e q u i p m e n t h e u s e dw h e n h e g o t t h e r e .

A l e x w a s a b o u t t o b e c o m e e m e r i t u s p r o f e s s o r o f p h y s io l o g y a t t h eH a r v a r d M e d i c a l S c h oo l w h e n w e m e t . H e l i v ed in t h e B l u e H i l ls s e c t io n o fM i l t on , a B o s t o n s u b u r b . A r o u n d 1 9 10 , a s a y o u n g f a c u l ty m e m b e r , h e r o d eto w o r k o n h o rs e b a c k , s t a b l i n g hi s a n i m a l d u r i n g w o r k i n g h o u r s i n a b a r no n H u n t i n g t o n A v e n u e n e a r t h e m e d i c a l sc ho ol. D u r i n g t h e w a r t i m e 1 9 40 s ,a s a m e m b e r o f a m a p p i n g e x p e d i t io n o r g a n i z e d b y t h e U . S. G e o l og ic a lS u r ve y , h e p i l o te d h i s o w n p l a n e w h i l e t a k i n g p i c t u r e s o v e r N o v a S c ot ia .

H e w as m id d le -ag ed w h en so meo n e d i s co v e red h o w to amp l i fy sma l l e lec-t r i c a l s ig n a l s u s in g th e v ac u u m tu b e , o n e o f t h e mo s t s ig n i f i c an t ev en t s i n t h eh i s to ry o f t ech n olo gy , an ad v an ce r a n k ed b y so me ev en h ig h e r t h a n th e mic ro -scope and te lescope in i t s impor tance to sc ience . Every d isc ip l ine f rom as t ron-o m y t h r o u g h z oo lo gy e n t e r e d i ts m o d e r n e r a a s s o on a s i ts m e a s u r i n g i n s t r u -m en t s i n c lu d ed e l ec t ro n ic c i rcu i t s t h a t c r ea t e l a rg e v o l t ag es o u t o f sma l l o n es .C e r t a i n l y n e u r o s c i e nc e w o u l d n o t b e w h a t w e k n o w w i t h o u t t h e v o l ta g e a m p l i-f ie rs in e lec t ron microscopes , computers , phys io log ica l s t imula to rs , and so on .

H a l D a v i s s t a t e s t h a t i n 1 9 2 3 A l e x " h a d a l r e a d y d e v e l o p e d a c a p a c i ty -c o u p l e d v a c u u m t u b e a m p l i f i e r t o i n c r e a s e t h e s e n s i t i v i t y o f h i s s t r i n g g a l -v a n o m e t e r , a n d w a s t h e f i rs t to e m p l o y a n a m p l i f i e r i n a p h y s io l o g ic a l ex p e r -im en t " (D av i s , 1 99 1 ). A r o u n d 1 9 3 0, w h e n A lex d ec id ed to m o d e rn ize h i s sy s -t e m , h is o p t io n s w e r e a n o t h e r s t r i n g g a l v a n o m e t e r o r t h e n e w v a c u u m t u b ea m p l i f i e r - p l u s - c a t h o d e - r a y - o s c i l l o s c o p e s y s t e m b e i n g u s e d b y a d v e n t u r o u sn e u r o p h y s i o l o g i s t s l i k e G a s s e r a n d E r l a n g e r i n S t . L o u i s . A c c o r d i n g t oD a v i s , t h e d e f i ci e nc i es o f t h e t h e n - a v a i l a b l e c a t h o d e r a y t u b e , w h o s e m o v i n gs p o t o f l i g h t c o u ld b e s e e n o n l y b y a p a r t i a l l y d a r k - a d a p t e d e y e, le d F o r b e st o s e l e c t t h e s t r i n g g a l v a n o m e t e r , b u t D o n L i n d s l e y h a s t o l d m e i t h a d n oa m p l i f i e r w h e n F o r b e s u s e d i t i n 1 9 3 3 . W h e n I a r r i v e d f i v e y e a r s l a t e r , as t r i n g g a l v a n o m e t e r w a s n o w h e r e t o b e s e e n in t h e H a r v a r d l a bo r at o ry .


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190 Robert GalambosI n 1 9 3 9, w h e n A l ex a d d e d m y n a m e t o t h e r e p o r t t h a t i n t r o d u c e d m et o g la s s m i c ro e l e ct ro d e s , A l b e r t G r a s s h a d o n l y r e c e n t l y c o m p l e t e d o u r

s t i m u l u s a n d r e c o r d i n g s y s t e m s , a " t h y r a t r o n s e t s i m i l a r t o t h e o n e u s e db y R e n s h a w " a n d " a c a p a c it y - c o u p le d p u s h - p u l l a m p l i f ie r c o n n e c te d w i t ha c a t h o d e - r a y o s c i l l o g r a p h " ( T h e r m a n e t a l . , 1 9 4 1 ) . I m e a s u r e d t h e b a tc o c h l e a r p o t e n t i a l s w i t h t h i s a m p l if ie r , a n d w i t h i n a w e e k i t w a s c l e a r t h eb a t e a r g e n e r a t e d f r e q u e n ci e s w e l l a bo v e t h e u p p e r l i m i t of A l b e r t' s a m p l i -f ie r r e sp o n s e . A f ew d a y s l a t e r , a s I w a s m o v i n g t h e b a t e x p e r i m e n t t o t h eC r u f t P h y s i c s l a b o r a t o r y a n d G .W . P i e rc e 's u n i q u e h i g h - f r e q u e n c y s y s t e m( N o y e s a n d P i e r c e , 1 9 38 ), A l b e r t t o ld m e h e f e l t b e t r a y e d . H e h a d a s k e dD a v i s a n d F o r b e s w h a t t h e u p p e r f r e q u e n c y l i m i t o f t h e n e w a m p l i f ie rs h o u l d b e , a n d w h e n t h e y s a i d 2 0 , 0 0 0 c y c l e s p e r s e c o n d h e k n e w t h e yw o u l d s h o r t l y w a n t m o r e , s o h e a r b i t r a r i l y r a i s e d t h e u p p e r l i m i t t o4 0 , 0 0 0 , w h i c h , a s t h e b a t s r e v e a l e d , w a s s t i l l n o t e n o u g h .W h a t a b o u t f u n d i n g ? W h o p a i d f o r s a la r i e s , s u p p l ie s , o v e r h e a d ? A l e xb o u g h t h i s o w n e q u i p m e n t a n d s u p p li es , a n d d o n a t e d h i s $ 6 00 y e a r l y s a l a rya l o ng w i t h e v e n m o r e p r i n c e ly s u m s t o t h e d e p a r t m e n t a n o n ym o u s l y. T h ew o r d o v e r h e a d e n t e r e d m y v o c a b u l a r y in t h e l a t e 1 9 4 0 s, a t w h i c h t im e u n i -v e r s i t i e s c o n s i d e r e d o n e p e r c e n t a w e l c o m e b o n a n z a . I n t h e m i d - 1 9 5 0 s ,w h e n I w a s d o i n g m y d u t y o n s t u d y s e ct io n s , I s o m e t i m e s s a w t h e s a m e p r o-p o s a l tw i c e , on c e a t a m e e t i n g o f t h e a g e n c y t h a t p a i d o v e r h e a d o n s a l a r i e so nly , a n d a g a i n a t t h e m e e t i n g o f t h e a g e n c y c a l c u l a t in g i t o n e q u i p m e n t a n ds u p p l i e s o n l y . D i s h o n e s t p e o p l e t u r n u p e v e r y w h e r e , b u t i n a l o n g c a r e e r Ih a v e a c t u a l l y k n o w n o n l y t w o c ro o k s w h o i n v e n t e d t h e i r d a t a .A l e x F o r b e s w a s a p i o n e e r A m e r i c a n e l e c t r o p h y s i o lo g i s t ; l i k e m e , h el o v e d t h e l a b o r a t o r y a n d c o n t i n u e d w o r k i n g i n o n e l o n g a f t e r o f f i c i a lr e t i r e m e n t . W a l la c e F e n n ' s s u m m a r y o f t h is g e n t l e m a n ' s m a n y c o n tr ib u -t i o n s i s a b e a u t i f u l t r i b u t e ( s e e i t i n t h e National Academy of SciencesMemoirs, Vol. 40) .Hallowell Dav is -L ou d Sounds and Hearing LossI n 1 9 4 2, j u s t a f t e r t h e P e a r l H a r b o r a t t a c k , H a l D a v i s w a s o f fe r ed t h e fo l-l o w in g a s s i g n m e n t : f in d o u t ho w m u c h a n d w h a t k i n d o f s o u n d i t t a k e s t oi n j u r e o r i n c a p a c i t a t e a m a n . A l i f e ti m e c o n s c i e n t i o u s o b j ec to r, h e r e s i g n e dh i s m e m b e r s h i p i n th e S o ci et y o f F r i e n d s a n d a c c ep t ed t h e a s s i g n m e n t(D av i s , 1991 p . 12 ). H a l co l l e c t ed t he fou r o f u s l i s t e d a s co -a u tho r s o f h i s1 9 5 0 m o n o g r a p h " T e m p o r a r y d e a f n e s s f o l l o w i n g e x p o s u r e t o l o u d t o n e sand no i s e " (D av i s e t a l . , 1950 ) , and w e p roceeded t o expose ou r e a r s t o t hes o u n d w a v e s e m i t t e d b y a s o -c a ll ed b u l lh o r n , t h e k i n d o f l o u d s p e a k e r t h eN a v y u s e d t o d e li v e r m e s s a g e s t o p e r s o n n e l w e a r i n g e a r p l u g s o n t h e b u s yf l ig h t d e c k o f a n a i r c r a f t c a rr ie r . W e s y s t e m a t i c a l l y v a r i e d t h e t h r e e s o u n dv a r i a b l e s - - i n t e n s i t y , f r e q u e n c y , a n d d u r a t i o n - - p r o d u c i n g i n o u r s e l v e si n c r e a s i n g l y l a r g e r t e m p o r a r y h e a r i n g l o s s e s , u n t i l w e n e a r e d c o m b i n a -


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Robert Galambos 191t io n s w e t h o u g h t m i g h t c a u s e a p e r m a n e n t l o ss . A t t h e e n d o f t h e p ro j ec t,H a l d e c i d e d t o f i n d o u t i f o u r p r e d i c t i o n s w e r e c o r r e c t , a n d t o ld u s t oe x p o s e h i s r i g h t e a r - - w e a l w a y s p r o t e c t e d h i s l e f t e a r - - t o a w i d e b a n dn o i s e a t 1 3 0 d B f o r 3 2 m i n u t e s . A s p r e d i c t e d , t h i s e x p o s u r e p e r m a n e n t l ys li c ed a f e w h u n d r e d H z o f f t h e h i g h e n d o f h i s e x i s t i n g c o n g e n i t a l h e a r -i n g l os s i n t h e 3 5 0 0 - 3 8 00 H z r eg i on . T h e m o n o g r a p h t h a t s u m m a r i z e s t h i sw o r k i s s t i l l q u o t e d i n t h e l i t e r a t u r e .

H a l b e g a n w e a r i n g h e a r i n g a i d s i n 1 9 7 9 , I i n 1 9 8 5 . W e a g r e e d t h o s ew a r t i m e e x p o s u r e s h a d n o t h i n g t o d o w i t h o u r p r e s b y c o u s i s . M y e v i d e n c es e e m s p a r t i c u l a r l y s t r o n g : w e e x p o s ed o n ly m y l e ft ea r , b u t m y m e a s u r e d l o ss -e s h a v e a l w a y s b e e n s y m m e t r i c a l , a n d I i n v a r i a b l y p u t t h e t e l e p h o n e t o m yle f t e a r , t h e o n e th a t t o o k a l l t h e b ea t in g , b eca u se I " h ea r b e t t e r " o n th a t s id e .

H a l c a l l e d h i s l a s t r e s e a r c h p r o j e c t " O l d T i m e E a r s . " I n 1 9 9 0 , h e c o n -v i n c ed 1 5 a g i n g h e a r i n g s p e c ia l is t s to j o in h i m i n s y s t e m a t i c a l l y d o c u m e n t -i n g t h e p r o g r e s s o f t h e i r h e a r i n g l o ss e s b y a ll a v a i l a b le t e s t s , a n d r e c r u i t e dC h a r l e s I . B e r l i n a n d L i n d a H o o d a t t h e K r e s g e H e a r i n g R e s e a r c h I n s t i t u t eo f t h e S o u t h i n N e w O r l e a n s t o a d m i n i s t e r t h e m . I n l a t e 1 99 5, a ll o f u se x c e p t H a l h a d o u r h e a r i n g t e s t e d o n c e a g a i n i n N e w O r l e a n s . H a l d i s -c h a r g e d h i s f i n a l o b l i g a t i o n t o t h e p r o j e c t i n 1 9 9 2 w h e n h i s t e m p o r a l b o n e sr e a c h e d t h e T e m p o r a l B o n e B a n k i n B o s t o n f o r h i s t o l o g i c a l a n a l y s i s .The Origins of Neuroscience-Clifford T Morgan and F. O. Schmit tM o r g a n is o n e o f t h e c o - a u t h o rs o f t h e D a v i s t e m p o r a r y h e a r i n g l os s m o n o-g r a p h . I n t h e s u m m e r o f 1 9 4 2, H a l s e n t t h e t w o o f u s t o W o o ds H o l e t o f i n do u t w h e t h e r u n d e r w a t e r e x p l os io n s a r e h a z a r d o u s f or t h e e a r s . S o m e p h y si -c i s t s w e r e e x p l o d i n g b o m b s i n t h e h a r b o r t h e r e , a n d w e w e r e s u p p o s e d t oj u m p i n a n d h a v e o u r h e a d s s u b m e r g e d w h e n t h is h a p p e n e d . W e s p e n t s ev -e r a l b e a u t if u l s u m m e r d a y s t a k i n g t u r n s j u m p i n g o ff t h e p i e r a t t h eO c e a n o g r a p h i c I n s t it u t e . T h e p l a n r e q u i r e d c o m p a r i n g b ef o re a n d a f t e ra u d i o g r a m s , a n d w e b e g a n w i t h b l a s t i n g c a p s d e t o n a t e d a t 5 0 f e e t o r s o .W h e n w e d e t e c t e d n o l o s s e s f o l l o w i n g d e t o n a t i o n s s o c l o s e t h a t w e w e r ea f ra i d w e m i g h t b e w o u n d e d b y s h r a p n el , w e b e g a n j u m p i n g i n w h e n t h eb l a s t e r s s i g n a l l e d a b o m b o f t h e i r s w a s a b o u t t o go o f f. T h e y s u p p l i e d u sw i t h p r e s s u r e d a t a f r o m t h e i r s e n s or s , a n d I r e c al l r e a l l y i m p r e s s i v e s h o c kw a v e s c o m p r e s s i n g m y b od y, b u t n e i t h e r o f u s e v e r r e c o rd e d a h e a r i n g lo ss .

M o r g a n c a m e t o H a r v a r d w i t h h i s n e w p s y c h ol o gy Ph . D . f r omR o c h e s t e r U n i v e r s i t y to w o r k w i t h K a r l L a s h le y , b u t b e f o re l o n g h e w a st r a v e l i n g t h r o u g h o u t t h e c o u n t r y fo r t h e N a t i o n a l D e f e n s e R e s e a r c hC o u n c i l h e l p i n g c o o r d i n a t e t h e e f fo r t s o f d i f f e r e n t l a b o r a t o r i e s w o r k i n g o nt h e s a m e o r s i m i l a r w a r t i m e p r o b le m s . W e w e r e c lo se p e r s o n a l f r ie n d s a n dl a b o r a t o r y c o l l e a g u e s . M o r g a n ' s P h . D . t h e s i s h a d s h o w n c e r t a i n b e h a v -i o r a l s e i z u r e s i n r a t s t o b e a u d i o g e n i c , n o t th e p r o d u c t o f f r u s t r a t i o n o ra n x i e t y a s N . R .F . M a i e r h a d c l a im e d ; t w o o f o u r j o i n t p a p e r s u s e d t h e b u l l-


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192 Robert G alambosh o r n , o u t s i d e o f o ff ic ia l h o u r s , t o c o n fi r m a n d e x p a n d t h i s p o i n t ( M o r g a na n d G a l a m b o s , 1 9 4 2 , 1 9 4 3 ) . W e a l s o m a d e p i t c h a n d l o u d n e s s m e a s u r e -m e n t s i n m a n ( M o r g a n e t a l., 1 9 5 1) , a n d i n 1 9 5 7 -5 8 w o r k e d o u t a lo n g a n dd i ff ic u l t c h a p t e r o n t h e n e u r a l b a s i s o f l e a r n i n g f o r t h e f i r s t Handbook o fNeurophysiology ( G a l a m b o s a n d M o r g a n , 1 9 6 0 ) .C l if f w e n t to Johns H opkins to cha i r it s Psycho logy dep ar tm en t in 1947and res igned in 1958 w hen he could no longer to l e ra t e the t ed ium of adm in-is t ra t ion . A few years la ter , royal t ies f rom his Introduction to Psychology(1961) m a de h im r ich . H e becam e pe r ipa tet i c , and se rved w i thou t p ay on thepsycho logy facu l ti e s a t t he U n ive r s i ty o f W iscons in a nd the U n ive r s i ty o fC a l i fo rn ia a t Sa n ta B arb a ra . In A us t in , T exas he w as loose ly a s soc ia t ed w i ththe Un ivers i ty of Texas , he lped foun d the Psychonom ic Socie ty , na m ed i t, andes tab l i shed and ed i t ed i t s j ou rn a l un t i l h i s un t im e ly dea th the re in 1976.H is Physiological Psychology, w r i t t e n i n s p a r e t i m e d u r i n g h i s w a rw o r k , w a s p u b l i s h e d i n 1 9 4 3 . I n it h e s a y s , " th e p r i m a r y g o a l o f p h y s i o -l o gi ca l p sy c h o l o g y is to e s t a b l i s h t h e p h y s i o lo g i c a l m e c h a n i s m s o f n o r m a lh u m a n a n d a n i m a l b e h a v i o r " ( M o r g a n 1 9 4 3 , p v i i ) . I t s 2 6 c h a p t e r s c o v e r ,i n s o m e 6 0 0 p a g e s , n o t h i n g b u t , a n d e s s e n t i a l l y e v e r y t h i n g k n o w n t h e na b o u t , w h a t w e c a l l n e u r o s c i e n c e t o d a y . T h e f o l l o w i n g p a r a g r a p h c o m e sf rom the in t roduc t ion to h i s t h i rd , 1965 , ed i t ion :

Perhaps no sub jec t d raw s upon so m any d i f f e ren t sc i ences andthe i r m ethod s as does phys io logica l psychology. E ver y sor t of pu reand app l i ed sc i en t i s t - -m a them at i c i an , phys ic i s t , chem is t , phys i -o logist , pharm acologis t , a nato m is t , ne urologis t , psychia t r i s t , e lec-t r i ca l eng inee r , a s w e l l a s psycho log i s t - -has been t ak ing pa r t i nour subjec t in one way or another (Morgan, 1965, p 9) .

I t c a n b e a r g u e d , a n d I d o , t h a t w h e n F r a n k S c h m i t t t h r e e y e a r s e a r -l i e r c o i n e d t h e w o r d " n e u r o s c i e n c e , " h e m e r e l y r e n a m e d a n e x i s t i n g d i s -c i p l i n e h a r d a t w o r k d o i n g e x a c t l y w h a t h e h a d i n m i n d ( t h e f i r s tPhysiologische Psychologie w a s p u b l i s h e d b y W i l h e lm W u n d t i n 1 8 7 3) .S c h m i t t' s e a r l y N e u r o s c i e n c e R e s e a r c h P r o g r a m A s s o c i at e s, o f w h o m Iw a s o n e , a r e a l l s p ec if ic e x a m p l e s o f t h e p h y s i c i s t s , c h e m i s t s , a n d b io lo -g i s t s o n M o r g a n ' s l is t ( S c h m i t t , 1 9 90 , p. 2 1 8) . F r a n k a n d C l if f l o o k e d a tt h e s a m e t h i n g t h r o u g h d i f fe r e n t g o g g le s . I c a n i m a g i n e C l if f c o n g r a t u -l a t i n g F r a n k o n h a v i n g r e c r u i t e d a ll t h o s e N o b e l P r i z e w i n n e r s t o j o i n th eo r d i n a r y b i o l o g i s t s , c h e m i s t s , a n d p h y s i c i s t s a l r e a d y t r y i n g t h e i r b e s t t od e s c r ib e t h e b r a i n c o r r e l a te s o f l e a r n i n g , m e m o r y, t h i n k i n g , m o t i v a t i o n ,a n d s o o n. O f c o u r se , t h is t a k e s a w a y n o t h i n g f ro m F r a n k S c h m i t t' s c o n-t r i b u t i o n t o t h e e f f o r t ; t h i s r e m a r k a b l e m a n o r g a n i z e d , p r o m o t e d , a n dc a t a l y z e d m u c h o f w h a t s u b s e q u e n t l y t r a n s p i r e d . B u t l e t h i s t o r y n o t e h ew a s n o t , a s s o m e c l a i m , t h e f i r s t t o d i s c o v e r t h e n e e d f o r e x t e n s i v e i n t e r -d i s c i p l i n a r y c o l l a b o r a t i o n s .


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Robert Galambos 193By 1962 , wh en F ran k Sch m i t t i nv i t ed me to j o in h is abou t - t o -be -o rga -

n i z ed N e u r o s c ie n c e R e s e a r c h P r o g r a m ( N R P) , I h a d s p e n t 1 0 y e a r s a s a m e m -be r o f Dave R ioch 's Wa l t e r Reed g roup , a h i s to r i ca l ly imp or t an t p ro to type o ft h e m o d e r n n e u r o s c i e n c e l a b o r a t o r y w h e r e d e p a r t m e n t l i n e s w e r e d e l i b e r -a t e ly b lu r r ed , and c ro ss -d is c ip l ine t h ink ing , t he ha l lm ark o f phys io log ica lp sycho log is t and n eu rosc i en t i s t a li ke , wa s t he ru l e. Fu r the rm ore , I had founda s imi l a r sp i r i t o f i n t e rd i s c ip l i na ry i n t e r ac t i on t o be t he wa y o f l if e in t h eM a g o u n g r o u p i n L o s A n g e l es w h e r e I s p e n t t h e s u m m e r o f 1 95 5.

N e u r o s c i e n c e a t t h e c o n c e p t u a l , t e x t b o o k , a n d l a b o r a t o r y l e v e l s m a yn o t h a v e b e e n n e w i n 1 9 6 2, b u t F r a n k ' s N R P c e r t a i n l y w a s . I t s fa c ul ty , th eA s s o c i a t e s , s p o k e o f t e n a n d e l o q u e n t l y f r o m t h e p l a t f o r m s h e c r e a t e d f o rt h e m . T h e e l e c t r o n m i c r o s c o p e h a d j u s t c o m e o f a g e ; t h e m o l e c u l a r bi ol o-g y r e v o lu t io n w a s b a r e l y u n d e r w a y ; n e u r o c h e m i s t r y w a s a t i ts t h r e s h o l do f u n p r e c e d e n t e d g r o w t h ; a n d t h e f i rs t c o g n it iv e e v o k e d p o t e n t i a l s h a dj u s t b e e n a v e r a g e d b y c o m p u t e r s . N o t h i n g l i k e t h i s h a d e v e r h a p p e n e db e fo r e, a n d t h e A s s o c ia t e s t ol d e a c h o t h e r a t W o r k S e s s io n s a n d A n n u a lM e e t in g s h o w t h e n e w m e t h o d s a n d d a t a w e r e t r a n s f o r m i n g o ld co n c ep tsa n d c r e a t i n g n e w o n e s . E a c h w a s a w o r l d - c l a s s e x p e r t i n h i s f i e l d , a n d t h ea u t h o r i t y a n d e l e g a n c e o f t h e i r p r e s e n t a t i o n s m a d e f o r m e m o r a b l e l e a r n -i n g e x p e r i e n c e s .T h e o r i g i n s a n d g o a l s o f S c h m i t t ' s N R P c a n b e t r a c e d d i r e c t l y t o h i se a r l i e r r e s p o n s e , i n t h e m i d - 19 5 0 s , to t h e 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 ha u t h o r i t i e s w h o a s k e d h i m " W h a t is b i o p h ys i cs ? " H e a n s w e r e d , i n 1 9 5 8, b yo r g a n i z i n g a m o n t h - l o n g " I n t e n s i v e S t u d y P r o g r a m " ( I S P ) i n B o u l d e r ,C o l o r a d o , a t w h i c h 6 1 e x p e r t s d e l i v e r e d l e c t u r e s w h i c h w e r e p u b l i s h e d i n1959 a s t he Biophysical Sciences-A Study Program. T h i s b o o k d e f i n e dt h e f i eld fo r t h e f i r s t ti m e a n d w a s i n s t r u m e n t a l i n th e c r e a t i o n o f t h eB iophys i ca l Soc i e ty .

A f ew y e a r s l a te r , S c h m i t t f o u n d h i m s e l f " in t e r e s t e d i n t h e p o s s ib i li tyt h a t i n f o r m a t i o n m i g h t b e t r a n s f e r r e d i n t h e b r a i n a n d c e n t r a l n e r v o u ss y s t e m n o t o n l y b y e l e ct r ic a l a c t io n w a v e s a l o n g n e u r a l n e t s , b u t a ls o b yf a s t t r a n s p o r t , p o s s ib l y t h r o u g h e x t r a c e l l u l a r s u b s t a n c e s " ( S c h m i t t , 1 9 9 0,p . 2 0 1 ) . I n o r d e r t o o r g a n i z e t h e e f f o r t t o f i n d o u t w h e t h e r t h e b r a i n a c t u -a l ly d o es w o r k t h i s w a y, h e s i m p l y e l a b o r a t e d a n d e x t e n d e d t h e p ro c e-d u r e s t h a t h a d s o s u c c e s s f u l l y s e t t l e d t h e q u e s t i o n , " W h a t i s b i o p h y s i c s ? "H e c o n c e i v e d , o r g a n i z e d , a n d f u n d e d w h a t c a m e t o b e c a l l e d t h eN e u r o s c i e n c e R e s e a r c h P r o g r a m . H e s e le c t ed e x p e r t s , t h e A s s o ci a te s , toa d v i s e h i m o n h o w t o p r o c ee d , a s s e m b l e d a s ta ff , a n d i n s t a l l e d i t in e x ce l-l e n t q u a r t e r s . B e c a u s e " f a s t t r a n s p o r t " w a s p r o m i n e n t i n h i s h y p o t h e s i s ,h i s o r i g i n a l 2 7 A s s o c i a t e s i n c l u d e d m a n y w i t h s p e c i a l k n o w l e d g e o f, o ri n t e r e s t i n , th e f a s t t r a n s f e r o f e l e m e n t a r y p a r t i c l e s ( e le c tr o n s a n d p ro -t o n s ) i n s ol id s a n d w a t e r s o l u t i o n s ; fi ve of t h e m w e r e p u r e p h y s i c a lc h e m i s t s , a n d f u l l y t w o - t h i r d s w e r e p r i m a r i l y p h y s i c i s t s o r c h e m i s t s . H ea ls o b e g a n p l a n n i n g a m o n t h - l o n g n e u r o s c i e n c e I S P a t B o u l d e r a n d c o n-


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194 Rober t Ga lambosvened it in 1966. This time he raised the number of experts delivering lec-tures to 65. Their contributions appeared a year later in what has beencalled the bible of Neuroscience, the first of the four volumes of T heNeurosciences: A S tudy Program. Interestingly, only about a third of thebook deals with the particular molecular biology questions that initiallyattracted Schmitt to the field.

The four S t u d y Pr o g r a m volumes received world-wide acclaim asauthor ita tive definitions and periodic updates of the field of neuroscience.Schmitt's NRP will also be remembered for its N R P Bu l l e ti n s, which wereconceived by Ted Melnechuk, an interdisciplinary writer who joined thestaff in 1963 as director of publications to help plan the Boulder ISP. Tedimmediately suggested that the Associates pinpoint the new findings andideas that might become topics on the Boulder program; then invite adozen world-class experts to a Work Session where one of the topics wouldbe discussed; and then prepare and dissemina te an edited version of theirdeliberations and conclusions. His ideas were accepted, and six such WorkSessions per year were promptly authorized; the first ones covered suchneuroscientific vanguards as biomolecular information storage, thesynapse, cell membranes, glial cells, brain correlates of memory, mathe-matical concepts of CNS function, and immunoneurology (a word, like"neuroscience" itself, first promulgated in the N RP Bu l l e t i n ) . Between1963 and 1972 the Bul let ins clarified the conceptual and empirical stateof research in 75 such neuroscience subfields. The Bul let ins became verypopular, and reached thousands of practicing and potentia l neuroscien-tists and science libraries around the world (few know about the two-dayWork Session on Extrasensory Perception I attended in the early days ofthe NRP; a Bul let in reporting it out was considered but rejected. Frankwould try almost anything in his search for enlightenment).

During my 20-years as an NRP Associate I attended all four Bouldermeetings and coauthored three of the Bul let ins , all made possible byFrank's vision, hard work and extraordinary executive abilities.Medical School and Mi l i tary ServiceMy best friend, when I was 10 years old, was named Wilfred Earl Allyn,Jr. His father was a doctor, and we occasionally snuck into his homelibrary to look at the pictures in his books. It was during this period of mylife that I first wanted to be a doctor. Later, after reading Paul DeKruif'sMicrobe Hunters , I had to be.At Oberlin I was a premed major, but on graduation, in 1935, in themiddle of the depression, financ ing a medical school education was out ofthe question. But the yearning would not go away, and finally, in 1942, mywife Jeannette and I decided it was now or never. Obviously the dreamcould come true only if she wen t to work to support three of us, which she


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R o b e r t G a l a m b o s 195d id . I w a s a c c e p t e d a t t h e U n i v e r s i t y o f R o c h e s t e r S c h oo l o f M e d i c i n e.W o r l d W a r I I w a s o n, a n d I en l i s te d a s a F i r s t L i e u t e n a n t i n t h e A r m yM e d i c a l S e r v i c e C o r p s . T h e w a r a n d m y m e d i c a l s c h o o l e d u c a t i o n e n d e da t a l m o s t t h e s a m e t i m e w i t h o u t m y e v e r s e r v i n g a d a y i n u n i f o r m .

B u t t h e r e i s m o r e t o t h i s m i l i t a r y h i s t o r y . I n 1 9 5 2 , d u r i n g t h e K o r e a nW a r , t h e d r a f t b o a r d c a l l e d m y n u m b e r . I r e p o r t e d f o r t h e p h y s i c a l e x a m -i n a t i o n , p a s s e d i t , a n d p r e p a r e d t o r e c e i v e m a r c h i n g o r d e r s . T h e s e n e v e rc a m e , a n d I l a t e r f o u n d o u t w h y . T h e A r m y h a d n e g l e c t e d t o d i s c h a r g e m ef r o m t h e M e d i c a l S e r v ic e C o r p s, w h i c h m e a n t I h a d t e c h n i c a l l y b e e n a so l-d i e r f or m o r e t h a n 10 y e a r s. T h e a u t o m a t i c a d v a n c e s i n r a n k a l o n g w i t ho t h e r p e r k s d u e m e w o u l d m e a n i n d u c t i n g m e a s p e r h a p s a L t . C o l o n e le n t i t l e d t o a b u n d l e o f a c c u m u l a t e d b a c k p ay , w h i c h m a d e s e n s e t o n o on e.A t R o c h e s t e r I w a s i n v o l v e d i n s e v e r a l e x p e r i m e n t s , o f w h i c h o n l y o n er e a c h e d p u b l i c a t i o n ( F e n n e t a l . , 1 9 4 9 ) . O t h e r e x p e r i m e n t s i n c l u d e dm i c r o e l e c t r o d e p e n e t r a t i o n s o f t h e c a t o p t ic n e r v e w i t h K a r l L o w y in t h ep s y c h o lo g y d e p a r t m e n t ; r e c t a l f e e d i n g o f p a r a l y z e d p o l i o m y e l i ti s p a t i e n t si n t h e i r o n lu n g ; a n d , w i t h J o s e B a r c h i l o n , t h e t r e a t m e n t o f a c u t e p o i so n -i ng b y t h e m u s h r o o m A m a n i t a p h a l l o i d e s .

I i n t e r n e d i n m e d i c in e a t E m o r y U n i v e r s i t y H o s p i t a l in A t l a n t a , a n df o r a n o t h e r y e a r d e b a t e d , w h i l e t e a c h i n g a n a t o m y t o m e d i c a l s t u d e n t st h e r e w i t h H a r l o w A d e s, w h e t h e r t o p r ac t ic e m e d i c in e o r r e t u r n t o t h e l a b-o r a t o r y . T h e l a b o r a t o r y w o n o u t , a n d I h a d t o c h o o s e b e t w e e n t h e W i l m e rI n s t i t u t e i n B a l t i m o r e a n d t h e P s y c h o a c o u s t i c L a b o r a t o r y ( P A L ) . T h e P A Lw a s S .S . ( S m i tt y ) S t e v e n s ' w a r t i m e l ab i n t h e b a s e m e n t o f H a r v a r d ' sM e m o r i a l H a l l , n o w n e w l y c i v i l i a n i z e d b u t s t i l l f u n d e d b y t h e O f f i c e o fN a v a l R e s e a r c h . W h e n I a s k e d S m i t t y w h y h e w a n t e d m e t o c om e , h e s a i dt h a t t h e w a r h a d c o n s u m e d a l l o u r b a s ic k n o w l e d g e a b o u t h e a r i n g , a n d w en e e d e d p u r e r e s e a r c h to g e n e r a t e m o r e b e f or e t h e f i g h t i n g b e g a n a g a i n .H a r v a r d I I , 1 9 4 7 - 5 1M y p l a n w a s s i m p le . T h e c a t s a n d I w o u l d c o n v e rs e , w i t h m e a s k i n g t h e q u e s -t io n s b y d e l i v e r in g c li ck s a n d t o n e s t o t h e i r e a r d r u m s , a n d t h e y r e p l y in g , o n eb ra in ce l l a t a t ime , t h ro u g h a mic ro e lec t ro d e . N o th eo ry , n o p reco n cep t io n s ;j u s t s i m p l e e x p e r i m e n t a l f a c t s . I a d o p t e d t h i s s t e r n p o s i t i o n b e c a u s e , a sr e c o u n t e d e l s e w h e r e ( G a l a m b o s , 1 9 9 2 a ) , H a l D a v i s a n d I h a d f o u n d i n h i b i -t i o n i n t h e a u d i t o r y n e r v e , a t o t a l l y u n e x p e c t e d e v e n t n e i t h e r t e a c h e r s n o rt e x tb o o k s h a d p r e p a r e d m e fo r. A p o x on b o t h t h e i r h o u s e s . T e a c h e r s a n db o o k s p ed d le d o g m a , t h e en e m y o f d is cov e ry , an d f ro m n o w o n I w o u ld b e l iev eonly w ha t I cou ld coax the ca ts to te ll me (ac tua l ly , as wi l l becom e c lear shor t -ly, mo s t o f o u r e l ec t ro d es h a d ce r t a in ly r e s t e d in th e co ch lea r n u c leu s , n o t t h en e r v e , a n d h a d I k n o w n t h i s t h e r e w o u l d h a v e b e e n n o r e a s o n f o r d i si ll us io n ).

A d o z e n p u b l i c a t i o n s c a m e o u t o f m y s e c o n d H a r v a r d p e r io d , o n e orm o r e w i t h c o l l a b o r a t o r s R e g B ro m i l ey , I r a H i r s h , J o h n R . H u g h e s , L a r r y


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196 Rober t Ga lambosK a h a n a , C l if f M o r g a n , J e r z y R o s e , W a l t e r R o s e n b l i t h , M a r kR o s e n z w e i g , a n d C a r r o l l L . W i l li a m s . O f t h e s e , t h e t h r e e w i t h J e r z yR o s e o n t h e m e d i a l g e n i c u l a t e c o n s u m e d t h e m o s t t i m e a n d e f f o r t . W em a p p e d t h e l o ca t io n of r e s p o n d i n g u n i t s i n t h a t n u c l e u s, a n d w h e t h e rt h e y r e s p o n d e d t o c l i c k s , n o i s e , o r t o n e s d e l i v e r e d m o n a u r a l l y a n d b i n -a u r a l l y . J e r z y l i k e d m y m i c r o e l e c t r o d e s a n d c a r r i e d s a m p l e s b a c k t oB a l t i m o r e s c o t c h - t a p e d i n s i d e t h e r e a r w i n d o w o f h i s c ar . V e r n o nM o u n t c a s t l e t o ld m e r e c e n t l y t h o s e h i g h i m p e d a n c e p i p e t t e s d id n o t w o r kw i t h t h e B a l t i m o r e l ow i n p u t - im p e d a n c e a m p l if ie r s, w h e r e u p o n J e r z yd e v i s e d t h e f a m o u s D o w b e n - R o s e m e t a l v e r s i o n a n d t h e J o h n s H o p k i n sl a b o r a t o r y e n t e r e d t h e s i n g l e u n i t b u s i n e s s .

I d id m o s t o f t h e w r i t in g o n t h e m e d i a l g e n i c u l a te p a p e r s , a n d w h e n w es e n t t h e m t o t h e e d i t o r i n 1 9 5 1 I t o l d J e r z y I w a s d e e p l y d i s a p p o i n t e d a th o w l i t t l e w e h a d l e a r n e d a f t e r s o m u c h e f f o r t . J e r z y , w h o h a d p r a c t i c e dp s y c h i a t r y i n t h e P a c i f i c d u r i n g W o r l d W a r I I , s o u g h t t o s o o t h e m e w i t ht h i s r e p l y : "M a y b e so , b u t t h e s e w i ll so o n b e t h e b e s t p a p e r s o n t h e m e d i a lg e n i c u l a t e e v e r p u b l i s h e d . " H e k n e w t h e y h a d t o b e , b e c a u s e f o r s e v e r a ly e a r s t h e r e w e r e n o o th e r s.C a t e x p e r i m e n t s w e r e a s m a l l fr a c ti o n o f w h a t w e n t o n a t P A L . E . G .B o r in g , th e d e p a r t m e n t c h a i r m a n , i n v i te d u s t o b r i n g o u r b r o w n b a g s a n dj o i n h i m a t l u n c h e v e r y d a y a r o u n d a h u g e o v a l t a b l e . G e o r g e A . M i l le r ,J . C . R . L i c k l i d e r , a n d I r a H i r s h , a m o n g o t h e r s , w e r e b e g i n n i n g t o b e c o m ef a m o u s . M y y o u n g e s t d a u g h t e r s p e n t h e r f i rs t y e a r i n th e S k i n n e r c ri bG e o r g e a n d I b u i l t , m o r e o r l e s s o v e r s e e n b y B . F . S k i n n e r h i m s e l f , i n t h el a b o r a t o r y s h o p . R u f u s G r a s o n a n d S t e v e S t a d l e r s o o n g r a d u a t e d f r o mt h a t s h o p t o f o rm t h e i r c o m p a n y t h a t s ol d t h e a m p l i f ie r s a n d a u d i o m e t e r st h e y h a d l e a r n e d t o pe rf ec t, a n d a lo n g w i t h a n o t h e r g r a d u a t e , R a l p hG e r b r a n d s , t h e f i r s t g e n e r a t i o n o f o p e r a n t c o n d i ti o n i n g t i m i n g a n d r e c o rd -i n g e q u i p m e n t . W a l t e r R o s e n b l i t h k e p t t a l k i n g a b o u t t h e N I H - f i n a n c e dc o m p u t e r b e i n g b u i l t n e a r b y , a t M I T ' s L i n c o ln L a b o r a t o r y , to p r o c e s s p h y s -i o l o g i c a l d a t a l i k e w h a t h e , M a r k R o s e n z w e i g , a n d I w e r e c o a x i n g o u t o fo u r c a t s , b u t t o m e t h e c o m p u t e r w a s a n u n n e c e s s a r y d i s t r a c t i o n . I w a ss t i l l t r y i n g t o f i n d t h e d a t a w o r t h p r o c e s s i n g .BekesyG e o r g v o n B e k e s y w a s b r o u g h t t o P A L i n 1 9 4 7 b y S m i t t y a n d E . B .N e w m a n f ro m S w e d e n , w h e r e h e h a d g o n e a f te r le a v i n g B u d a p e s t a t t h ee n d o f W o r l d W a r I I. W h e n I a r ri v e d , h e w a s s e t t i n g u p t o c o n t i n u e t h eb a s i l a r m e m b r a n e m e a s u r e m e n t s f o r w h i c h h e w o u l d r e c e i v e t h e N o b e lP r i z e . H e w a s a q u i e t m a n , a b a c h e l o r , w h o r a r e l y c o n t r i b u t e d t o t h ew o r d y i n t e r p l a y a t B o r i n g ' s t a b l e . H i s 8 3 - i t e m b i b l i o g r a p h y c i t e s o n l yt h r e e c o - a u t h o r e d p a p e r s . I r e m e m b e r h i m l a u g h i n g o n l y o n c e . W e w e r et a l k i n g w i t h a v i s it i n g s c i e n t is t f or w h o m I tr i e d t o e x p l a i n s o m e t h i n g i n


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R o b e r t G a l a m b o s 197m y h i g h -s c h o ol G e r m a n . I n o ti c ed B e k e s y l a u g h i n g , w i t h h i s h a n d h e l do v e r h i s m o u t h , a n d w h e n I a s k e d h i m l a t e r w h a t h a d b e e n f u n n y h e s a i dm y G e r m a n h a s a s t ro n g H u n g a r i a n a c ce n t (I l e a r n e d m y l im i t e d v oc a bu -l a r y o f H u n g a r i a n w o r d s a s a c h il d o v e r h e a r i n g c o n v e r s a t i o n s b e t w e e n m yp a r e n t s a n d o t h e r s ) .

D o n G r i f f i n s a y s t h e Y a l e b a t m a n , A l v i n N o v i c k , v i s i t e d B e k e s y i n1 9 5 3 o r 1 9 5 4 t o s e e k a d v i c e o n b a t h e a r i n g m a t t e r s b u t l e f t w i t h o u t a n y .B e k e s y w a s s k e p t i c a l a b o u t t h e w h o l e e c h o l o ca t i o n i d e a a n d s a i d t h e e m i t -t e d s o u n d s w e r e p r o b a b l y j u s t n o is e b u r s t s . A f ew y e a r s l a t e r a f t e r a t t e n d -i n g a s e m i n a r g i v e n b y a v i s i t i n g b a t m a n f r o m B r o w n U n i v e r s i t y , J i mS i m m o n s , B e k e s y w a s h e a r d t o sa y m a y b e t h e r e w a s s o m e t h i n g to t h ei d e a a f t e r a l l. B e k e s y a n d I s a w e a c h o t h e r a l m o s t d a i l y f or f o u r y e a r s , b u tw e n e v e r o n c e t a l k e d a b o u t b a t s . I s i t p o s s i b l e h e h a d n o t r e a d t h e b a tp a p e r s p u b l i s h e d 1 0 y e a r s e a r l ie r ?

A n o t h e r s t r a n g e t h i n g . I n 1 94 7 , I c a m e u p o n a b r i e f r e p o r t ( in aj o u r n a l I h a v e s in c e b e e n u n a b l e t o fi nd ) of m i c r o e l e c t r o d e e x p e r i m e n t sB e k e s y a n d a p e r s o n n a m e d H a m b u r g e r h a d d o ne on t h e c a t c o c h le a rn u c l e u s i n S w e d e n . T h e y c o n f i r m e d o u r 1 9 4 3 r e s u l t s a n d i n a d d i t i o nd e m o n s t r a t e d h i s t o lo g i c a l l y t h a t t h e i r e l e c t r o d e s h a d b e e n i n t h ec o c h l e a r n u c l e u s , n o t t h e a u d i t o r y n e r v e a s H a l D a v i s a n d I h a dc l a i m e d . O u r n o t e i n S c i e n c e s a y i n g w e h a d d i s c o v e r e d t h i s e m b a r -r a s s i n g f ac t o u r s e l v e s h a d j u s t a p p e a r e d ( G a l a m b o s a n d D a v i s , 19 48 ).W h e n I a s k e d B e k e s y w h y h e h a d n o t t o l d u s h e k n e w i t a l l a l o n g , h es a i d o u r e x p e r i m e n t a l f i n d i n g s h a d b e e n c o r r e ct l y r e p o r t e d , a n d h eb e l i e v e d o n e s h o u l d n o t e m p h a s i z e t h e m i s t a k e s i n a p u b l i c a t i o n u n l e s st h e y a l t e r t h e d a t a .

W e c o ll a b o r a te d i n o n ly o n e m e a s u r e m e n t . H i s q u e s t i o n w a s w h a t a ne a r d r u m l oo k s l ik e a s it r u p t u r e s . I e x p o s ed t h e e a r d r u m o f a n a n e s -t h e t i z e d g u i n e a p i g f r o m t h e i n s i d e b y r e m o v i n g t h e w a l l o f t h e b u l l a , a n dw e a d j u s t e d t h e l e n s o f a F a s t a x c a m e r a s o t h a t t h e e a r d r u m f il le d a3 5 m m m o v ie fi l m fr a m e . F a s t a x c a m e r a s c a n r u n t h o u s a n d s o f f r a m e sp a s t t h e l e n s e v e r y s ec o nd . B e k e s y f ix e d t h i n g s so t h a t t h e c a m e r a b e g a nr o l li n g a m o m e n t b e f o r e a s t a r t e r ' s p i s to l f ir e d a c a r t r i d g e n e x t t o t h e p i g 'se ar . E v e r y t h i n g w o r k e d . T h e e a r d r u m s h a t t e r s i n t o f r a g m e n t s t h a t f ly i na l l d i r e c t i o n s . T h e p i c t u r e s w e r e s p e c t a c u l a r , b u t I d o n ' t r e m e m b e r w h yB e k e s y w a n t e d t h e m o r w h a t h a s h a p p e n e d to t h e m .

O n e S u n d a y a f t e r n o o n I a c c o m p a n i e d h i m t o th e B o s t o n F i n e A r t sM u s e u m . H e h a d a n a p p o i n t m e n t w i t h t h e e g y p t o l o g i s t , w h o t o o k u s t o ab a s e m e n t s t o r a g e a r e a t o s ee th e i t e m s B e k e s y h a d i n m i n d . B e k e s y col-l e c t e d s u c h t h i n g s a n d w i l l e d t h e m a l l t o t h e N o b e l F o u n d a t i o n . H e t o l dm e t h a t w h e n h e r e c e i ve d h is P r i z e h e v i s it e d t h e K i n g o f S w e d e n i n h i so f f i c e , a s w a s c u s t o m a r y , a n d w h e n h e s a w a n E g y p t i a n a r t i f a c t o n t h es h e l f b e h i n d t h e K i n g's h e a d h e c o m m e n t e d o n i t, w h e r e u p o n t h e t w o o ft h e m s p e n t a n h o u r t a l k i n g a b o u t t h e h o b b y t h e y s h a r e d .


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198 Robert GalambosBekesy was also a historian-philosopher of science. For instance, he

classified experimental problems in the following concise and amusingway:

Problems arise in a variety of ways, and it is often worthwhileto list the forms that they may take. Thus we can distinguishthe following:1. The classical problem, which has had much effort expendedupon it, but without any acceptable solution.2. The premature problem, which often is poorly formulated,or is not susceptible to attack.3. The strategic problem, which seeks data on which a choice maybe made between two or more basic assumptions or principles.4. The stimula ting problem, which may lead to reexaminat ion ofaccepted principles and may open up new areas for exploration.5. The stati stical question, which may be only a survey of pos-sibilities.6. The unimportant problem, which is easy to formulate andeasy to solve.7. The embarrassing question, commonly arising at meetingsin discussion of a paper, and rarely serving any useful purpose.8. The pseudo problem, usually the consequence of differentdefinitions or metho

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Robert Galambos

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