Quantum Conciencia
5
7/27/2019 Quantum Conciencia http://slidepdf.com/reader/full/quantum-conciencia 1/5 Proc. Nadl. Acad. Sci. USA Vol. 89, pp. 11357-11361, December 1992 Biophysics Quantum aspects of brain activity and the role of consciousness FRIEDRICH BECK* AND JOHN C. ECCLESt *Institut for Kernphysik, Technische Hochschule Darmstadt, D-6100 Darmstadt, Germany; and tMax-Planck-Institut fOr Hirnforschung, D-6000 Frankfurt a.M. 71, Germany Contributed by John C. Eccles, July 21, 1992 ABSTRACT The relationship ofbrain activity to conscious intentions is considered on the basis of the functional micro- structure of the cerebral cortex. Each ming nerve impulse causes the emission of transmitter molecules by the process of exocytosis. Since exocytosis is a quantal phenomenon of the presynaptic vesiculargrid with a probability much less than 1, we present a quantum mechanical model for It based on a tunneling process of the trigger mechanism. Consciousness manifests itself in mental intentions. The consequent voluntary actions become effective by momentary increases of the prob- ability of vesicular emission in the thousands of synapses on each pyramidal cell by quantal selection. There has been an increasing interest in the relations of quantum mechanics, brain activities, and consciousness (1- 4). On the part of quantum physics the impetus camefrom the interpretation of the measuring process, which is still on debate, even after more than 60 years of overwhelmingly successful applications of quantum theory. However, there is the question from neuroscience (5) whether quantum action is needed to understand the functioning of the brain in its subtle relations to experience, memory, and consciousness. It was Wigner (6), in his stringentanalysis of the conse- quences of measurements in a Stern-Gerlach experiment, who first speculated that the von Neumann collapse of the wave function (7) actually occurs by an act of consciousness in the human brain and is not describable in terms of ordinary quantum mechanics. More recently similar ideas were put forward and partly combined with Everett s many-world interpretation (8) of quantum theory 2). Other authors (e.g., ref. 1) have related quantum theory to consciousness on the basis of the usual interpretation of the state vector as a superposition of actualities [or propensities, in Popper s nomenclature 9)]. Not much connection has been made, however, to the empirically established facts of brain phys- iology, nor have thoseauthors attempted to locate a quantal process in the functional microsites of the neocortex. In this work we contribute to filling this gap by putting forward a quantum mechanical description of bouton exocy- tosis. The next section gives an outline of the structure and activity of the neocortex as introduction to the quantum mechanical model. This is set up in the subsequent section. Finally, the hypothesis for a coherent coupling of the indi- vidual probability amplitudes is introduced. In this model action becomes possible by influence of a conscious will. Neocortical Activity Fig. 1A illustrates the universally accepted six laminaeof the neocortex (10) with two large pyramidal cells in lamina V, three in lamina III, and two in lamina II. The pyramidal apical dendrites finish in a tuft-like branching in lamina I (Fig. 2A). There is agreement by Peters and Fleischhauer and their associates (12, 13) that the apical bundles or clusters dia- grammatically shown in Fig. 2B are the basic anatomical units of the neocortex. They are observed in all areas of the cortex that have been investigated and in all mammals, including humanst. It has been proposed that these bundles are the cortical units for reception 5), which would give them a preeminent role. Since they are composed essentially of dendrites, the name dendron was adopted. Fig. 1B illustrates a typical spine synapse that makes an intimate contact with an apical dendrite of a pyramidal cell (Fig. 1; Fig. 3 A and D). The ultrastructure ofsuch a synapse has been intensively studied by Akert and his associates (14, 15). From the inner surface of a bouton confronting the synaptic cleft (d in Fig. 1B, the active site in Fig. 3A), dense projections in triangular array form the presynaptic vesicular grid (PVG) (Fig. 3 A-E). Fig. 3B is a photomicrograph of a tangential section of a PVG showing the dense projections in triangular array with the faint synaptic vesicles fitting snugly in hexagonal array. The spherical synaptic vesicles, 50-60 A in diameter, with their content of transmitter mol- ecules, can be seen in the idealized drawings of the PVG (Fig. 3 C and D) with the triangularly arranged dense projections, the active zone, and the hexagonal array of synaptic vesicles (14, 15). In a more primitive form the PVG can be seen in synapses of the fish Mauthner cell 17). A nerve impulse propagating into a bouton causes a process called exocytosis. At most a nerve impulse evokes a single exocytosis from a PVG (Fig. 3 D and G). This limitation is probably due to the vesicles being embedded in the paracrystalline PVG (Fig. 3 B-E). Exocytosis is the basic unitary activity of the cerebral cortex. Each all-or-nothing exocytosis of synaptic transmit- ter results in a brief excitatory postsynaptic depolarization (EPSP). Summation by electrotonic transmission of many hundreds of these milli-EPSPs is required for an EPSP large enough (10-20 m V) to generate the discharge ofan impulse by a pyramidal cell. This impulse will travel along its axon (Fig. 2A) to make effective excitation at its many synapses. This is the conventional macro-operation of a pyramidal cell of the neocortex (Fig. 1) and it can be satisfactorily described by classical physics and neuroscience, even in the most complex design of network theory and neuronal group se- lection (10, 18, 19). Exocytosis has been intensivelystudied in the mammalian central nervous system, where it is meanwhile possible to refine the study by utilizing a single excitatory impulse to generate EPSPs in single neurons that are being studied by intracellular recording. The initial studies were on the mono- synaptic action on motoneurons by single impulses in the large Ia afferent fibres from muscle (20). More recently (21) Abbreviations: EPSP, excitatory postsynaptic depolarization; PVG, presynaptic vesicular grid; SMA, supplementary motor area. lApproximate values can be given forthe synaptic connectivity of an apical bundle. The input would be largely by the spine synapses (Figs. 1 and 2A), of which there would be over 5000 on a lamina V apicaldendrite with its lateral branches and terminal tuft (Fig. 2A), but more usually there would be about 2000. If there are 70-100 apical dendrites in a bundle, the number of total spine synapses would be well over 100,000. 11357 The publication costs of this article were defrayed in part by page charge payment. This article must therefore be hereby marked advertisement in accordance with 18 U.S.C. §1734 solely to indicate this fact.
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Transcript of Quantum Conciencia
7/27/2019 Quantum Conciencia
http://slidepdf.com/reader/full/quantum-conciencia 1/5
P r o c . N a d l . A c a d . S c i . USAV o l . 8 9 , p p . 1 1 3 5 7 - 1 1 3 6 1 , D e c e m b e r 1 9 9 2B i o p h y s i c s
Quantum a s p e c t s o f b r a i n a c t i v i t y a n d t h e r o l e o f c o n s c i o u s n e s s
F R I E D R I C H B E C K * AND JOHN C . E C C L E S t
* I n s t i t u t f o r K e r n p h y s i k , T e c h n i s c h e H o c h s c h u l e D a r m s t a d t , D - 6 1 0 0 D a r m s t a d t , G e r m a n y ; a n d t M a x - P l a n c k - I n s t i t u t f O r H i r n f o r s c h u n g , D - 6 0 0 0 F r a n k f u r t a . M .7 1 , G e r m a n y
C o n t r i b u t e d b y J o h n C . E c c l e s , J u l y 2 1 , 1 9 9 2
ABSTRACT T h e r e l a t i o n s h i p o f b r a i n a c t i v i t y t o c o n s c i o u si n t e n t i o n s i s c o n s i d e r e d o n t h e b a s i s o f t h e f u n c t i o n a l m i c r o -
s t r u c t u r e o f t h e c e r e b r a l c o r t e x . E a c h m i n g n e r v e i m p u l s ec a u s e s t h e e m i s s i o n o f t r a n s m i t t e r m o l e c u l e s b y t h e p r o c e s s o fe x o c y t o s i s . S i n c e e x o c y t o s i s i s a q u a n t a l p h e n o m e n o n o f t h ep r e s y n a p t i c v e s i c u l a r g r i d w i t h a p r o b a b i l i t y m u c h l e s s t h a n 1 ,w e p r e s e n t a q u a n t u m m e c h a n i c a l m o d e l f o r I t b a s e d o n at u n n e l i n g p r o c e s s o f t h e t r i g g e r m e c h a n i s m . C o n s c i o u s n e s sm a n i f e s t s i t s e l f i n m e n t a l i n t e n t i o n s . T h e c o n s e q u e n t v o l u n t a r ya c t i o n s b e c o m e e f f e c t i v e b y m o m e n t a r y i n c r e a s e s o f t h e p r o b -a b i l i t y o f v e s i c u l a r e m i s s i o n i n t h e t h o u s a n d s o f s y n a p s e s o ne a c h p y r a m i d a l c e l l b y q u a n t a l s e l e c t i o n .
T h e r e h a s b e e n a n i n c r e a s i n g i n t e r e s t i n t h e r e l a t i o n s o fq u a n t u m m e c h a n i c s , b r a i n a c t i v i t i e s , a n d c o n s c i o u s n e s s ( 1 -4 ) . On t h e p a r t o f q u a n t u m p h y s i c s t h e i m p e t u s c a m e f r o m t h ei n t e r p r e t a t i o n o f t h e m e a s u r i n g p r o c e s s , w h i c h i s s t i l l o nd e b a t e , e v e n a f t e r m o r e t h a n 6 0 y e a r s o f o v e r w h e l m i n g l ys u c c e s s f u l a p p l i c a t i o n s o f q u a n t u m t h e o r y . H o w e v e r , t h e r e i st h e q u e s t i o n f r o m n e u r o s c i e n c e ( 5 ) w h e t h e r q u a n t u m a c t i o ni s n e e d e d t o u n d e r s t a n d t h e f u n c t i o n i n g o f t h e b r a i n i n i t ss u b t l e r e l a t i o n s t o e x p e r i e n c e , m e m o r y , a n d c o n s c i o u s n e s s .
I t w a s W i g n e r ( 6 ) , i n h i s s t r i n g e n t a n a l y s i s o f t h e c o n s e -q u e n c e s o f m e a s u r e m e n t s i n a S t e r n - G e r l a c h e x p e r i m e n t ,who f i r s t s p e c u l a t e d t h a t t h e v o n Neumann c o l l a p s e o f t h ew a v e f u n c t i o n ( 7 ) a c t u a l l y o c c u r s b y a n a c t o f c o n s c i o u s n e s si n t h e h u m a n b r a i n a n d i s n o t d e s c r i b a b l e i n t e r m s o f o r d i n a r yq u a n t u m m e c h a n i c s . M o r e r e c e n t l y s i m i l a r i d e a s w e r e p u t
f o r w a r d a n d p a r t l y c o m b i n e d w i t h E v e r e t t s m a n y - w o r l di n t e r p r e t a t i o n ( 8 ) o f q u a n t u m t h e o r y 2 ) . O t h e r a u t h o r s ( e . g . ,r e f . 1 ) h a v e r e l a t e d q u a n t u m t h e o r y t o c o n s c i o u s n e s s o n t h eb a s i s o f t h e u s u a l i n t e r p r e t a t i o n o f t h e s t a t e v e c t o r a s as u p e r p o s i t i o n o f a c t u a l i t i e s [ o r p r o p e n s i t i e s , i n P o p p e r sn o m e n c l a t u r e 9 ) ] . N o t m u c h c o n n e c t i o n h a s b e e n m a d e ,h o w e v e r , t o t h e e m p i r i c a l l y e s t a b l i s h e d f a c t s o f b r a i n p h y s -i o l o g y , n o r h a v e t h o s e a u t h o r s a t t e m p t e d t o l o c a t e a q u a n t a lp r o c e s s i n t h e f u n c t i o n a l m i c r o s i t e s o f t h e n e o c o r t e x .
I n t h i s w o r k we c o n t r i b u t e t o f i l l i n g t h i s g a p b y p u t t i n gf o r w a r d a q u a n t u m m e c h a n i c a l d e s c r i p t i o n o f b o u t o n e x o c y -t o s i s . T h e n e x t s e c t i o n g i v e s a n o u t l i n e o f t h e s t r u c t u r e a n da c t i v i t y o f t h e n e o c o r t e x a s i n t r o d u c t i o n t o t h e q u a n t u mm e c h a n i c a l m o d e l . T h i s i s s e t u p i n t h e s u b s e q u e n t s e c t i o n .F i n a l l y , t h e h y p o t h e s i s f o r a c o h e r e n t c o u p l i n g o f t h e i n d i -
v i d u a l p r o b a b i l i t y a m p l i t u d e s i s i n t r o d u c e d . I n t h i s m o d e la c t i o n b e c o m e s p o s s i b l e b y i n f l u e n c e o f a c o n s c i o u s w i l l .
N e o c o r t i c a l A c t i v i t y
F i g . 1A i l l u s t r a t e s t h e u n i v e r s a l l y a c c e p t e d s i x l a mi n a e o f t h en e o c o r t e x ( 1 0 ) w i t h t w o l a r g e p y r a m i d a l c e l l s i n l a m i n a V ,t h r e e i n l a m i n a I I I , a n d t w o i n l a m i n a I I . T h e p y r a m i d a l a p i c a ld e n d r i t e s f i n i s h i n a t u f t - l i k e b r a n c h i n g i n l a m i n a I ( F i g . 2 A ) .T h e r e i s a g r e e m e n t b y P e t e r s a n d F l e i s c h h a u e r a n d t h e i ra s s o c i a t e s ( 1 2 , 1 3 ) t h a t t h e a p i c a l b u n d l e s o r c l u s t e r s d i a -
g r a m m a t i c a l l y s h o w n i n F i g . 2B a r e t h e b a s i c a n a t o m i c a lu n i t s o f t h e n e o c o r t e x . T h e y a r e o b s e r v e d i n a l l a r e a s o f t h ec o r t e x t h a t h a v e b e e n i n v e s t i g a t e d a n d i n a l l m a m m a l s ,i n c l u d i n g h u m a n s t . I t h a s b e e n p r o p o s e d t h a t t h e s e b u n d l e sa r e t h e c o r t i c a l u n i t s f o r r e c e p t i o n 5 ) , w h i c h w o u l d g i v e t h e ma p r e e m i n e n t r o l e . S i n c e t h e y a r e c o m p o s e d e s s e n t i a l l y o fd e n d r i t e s , t h e name d e n d r o n w a s a d o p t e d .
F i g . 1 B i l l u s t r a t e s a t y p i c a l s p i n e s y n a p s e t h a t m a k e s a n
i n t i m a t e c o n t a c t w i t h a n a p i c a l d e n d r i t e o f a p y r a m i d a l c e l l( F i g . 1 ; F i g . 3 A a n d D ) . T h e u l t r a s t r u c t u r e o f s u c h a s y n a p s eh a s b e e n i n t e n s i v e l y s t u d i e d b y A k e rt a n d h i s a s s o c i a t e s ( 1 4 ,1 5 ) . F r o m t h e i n n e r s u r f a c e o f a b o u t o n c o n f r o n t i n g t h e
s y n a p t i cc l e f t
( d i n F i g . 1 B , t h e a c t i v e s i t e i n F i g . 3 A ) , d e n s ep r o j e c t i o n s i n t r i a n g u l a r a r r a y f o r m t h e p r e s y n a p t i c v e s i c u l a rg r i d ( P V G ) ( F i g . 3 A - E ) . F i g . 3 B i s a p h o t o m i c r o g r a p h o fa t a n g e n t i a l s e c t i o n o f a PVG s h o w i n g t h e d e n s e p r o j e c t i o n si n t r i a n g u l a r a r r a y w i t h t h e f a i n t s y n a p t i c v e s i c l e s f i t t i n gs n u g l y i n h e x a g o n a l a r r a y . T h e s p h e r i c a l s y n a p t i c v e s i c l e s ,5 0 - 6 0 A i n d i a m e t e r , w i t h t h e i r c o n t e n t o f t r a n s m i t t e r m o l -e c u l e s , c a n b e s e e n i n t h e i d e a l i z e d d r a w i n g s o f t h e PVG ( F i g .3 C a n d D ) w i t h t h e t r i a n g u l a r l y a r r a n g e d d e n s e p r o j e c t i o n s ,t h e a c t i v e z o n e , a n d t h e h e x a g o n a l a r r a y o f s y n a p t i c v e s i c l e s( 1 4 , 1 5 ) . I n a m o r e p r i m i t i v e f o r m t h e PVG c a n b e s e e n i ns y n a p s e s o f t h e f i s h M a u t h n e r c e l l 1 7 ) .A n e r v e i m p u l s e p r o p a g a t i n g i n t o a b o u t o n c a u s e s a
p r o c e s s c a l l e d e x o c y t o s i s . A t m o s t a n e r v e i m p u l s e e v o k e s as i n g l e e x o c y t o s i s f r o m a PVG ( F i g . 3 D a n d G ) . T h i sl i m i t a t i o n i s p r o b a b l y d u e t o t h e v e s i c l e s b e i n g e m b e d d e d i nt h e p a r a c r y s t a l l i n e PVG ( F i g . 3 B - E ) .
E x o c y t o s i s i s t h e b a s i c u n i t a r y a c t i v i t y o f t h e c e r e b r a lc o r t e x . E a c h a l l - o r - n o t h i n g e x o c y t o s i s o f s y n a p t i c t r a n s m i t -t e r r e s u l t s i n a b r i e f e x c i t a t o r y p o s t s y n a p t i c d e p o l a r i z a t i o n( E P S P ) . S u m m a t i o n b y e l e c t r o t o n i c t r a n s m i s s i o n o f manyh u n d r e d s o f t h e s e m i l l i - E P S P s i s r e q u i r e d f o r a n EPSP l a r g ee n o u g h ( 1 0 - 2 0 mV) t o g e n e r a t e t h e d i s c h a r g e o f a n i m p u l s eb y a p y r a m i d a l c e l l . T h i s i m p u l s e w i l l t r a v e l a l o n g i t s a x o n( F i g . 2 A ) t o m a k e e f f e c t i v e e x c i t a t i o n a t i t s many s y n a p s e s .T h i s i s t h e c o n v e n t i o n a l m a c r o - o p e r a t i o n o f a p y r a m i d a l c e l lo f t h e n e o c o r t e x ( F i g . 1 ) a n d i t c a n b e s a t i s f a c t o r i l y d e s c r i b e db y c l a s s i c a l p h y s i c s a n d n e u r o s c i e n c e , e v e n i n t h e m o s tc o m p l e x d e s i g n o f n e t w o r k t h e o r y a n d n e u r o n a l g r o u p s e -l e c t i o n ( 1 0 , 1 8 , 1 9 ) .
E x o c y t o s i s h a s b e e n i n t e n s i v e l y s t u d i e d i n t h e m a m m a l i a n
c e n t r a l n e r v o u s s y s t e m , w h e r e i t i s m e a n w h i l e p o s s i b l e t or e f i n e t h e s t u d y b y u t i l i z i n g a s i n g l e e x c i t a t o r y i m p u l s e t o
g e n e r a t e E P S P s i n s i n g l e n e u r o n s t h a t a r e b e i n g s t u d i e d b yi n t r a c e l l u l a r r e c o r d i n g . T h e i n i t i a l s t u d i e s w e r e o n t h e m o n o -s y n a p t i c a c t i o n o n m o t o n e u r o n s b y s i n g l e i m p u l s e s i n t h el a r g e I a a f f e r e n t f i b r e s f r o m m u s c l e ( 2 0 ) . M o r e r e c e n t l y ( 2 1 )
A b b r e v i a t i o n s : E P S P , e x c i t a t o r y p o s t s y n a p t i c d e p o l a r i z a t i o n ; P V G ,p r e s y n a p t i c v e s i c u l a r g r i d ; S M A , s u p p l e m e n t a r y m o t o r a r e a .l A p p r o x i m a t e v a l u e s c a n b e g i v e n f o r t h e s y n a p t i c c o n n e c t i v i t y o f a na p i c a l b u n d l e . T h e i n p u t w o u l d b e l a r g e l y b y t h e s p i n e s y n a p s e s( F i g s . 1 a n d 2 A ) , o f w h i c h t h e r e w o u l d b e o v e r 5 0 0 0 o n a l a m i n a Va p i c a l d e n d r i t e w i t h i t s l a t e r a l b r a n c h e s a n d t e r m i n a l t u f t ( F i g . 2 A ) ,b u t m o r e u s u a l l y t h e r e w o u l d b e a bo u t 2 0 0 0 . I f t h e r e a r e 7 0 - 1 0 0a p i c a l d e n d r i t e s i n a b u n d l e , t h e n u m b e r o f t o t a l s p i n e s y n a p s e sw o u l d b e w e l l o v e r 1 0 0 , 0 0 0 .
1 1 3 5 7
T h e p u b l i c a t i o n c o s t s o f t h i s a r t i c l e w e r e d e f r a y e d i n p a r t b y p a g e c h a r g ep a y m e n t . T h i s a r t i c l e m u s t t h e r e f o r e b e h e r e b y m a r k e d a d v e r t i s e m e n ti n a c c o r d a n c e w i t h 1 8 U . S . C . § 1 7 3 4 s o l e l y t o i n d i c a t e t h i s f a c t .
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1 1 3 5 8 B i o p h y s i c s : B e c k a n d E c c l e s
- 6 0 / i ml-a~M;-
F I G . 1 . ( A ) T h r e e - d i m e n s i o n a l c o n s t r u c t b y S z e n t a g o t h a i ( 1 0 )s h o w i n g c o r t i c a l n e u r o n s o f v a r i o u s t y p e s . ( B ) D e t a i l e d s t r u c t u r e o fa s p i n e ( s p ) s y n a p s e o n a d e n d r i t e ( d e n ) ; s t , a x o n t e r m i n a t i n g i ns y n a p t i c b o u t o n o r p r e s y n a p t i c t e r m i n a l ( p r e ) ; s v , s y n a p t i c v e s i c l e s ;c , p r e s y n a p t i c v e s i c u l a r g r i d ( P V G i n t e x t ) ; d , s y n a p t i c c l e f t ; e ,p o s t s y n a p t i c m e m b r a n e ; a , s p i n e a p p a r a t u s ; b , s p i n e s t a l k ; m ,m i t o c h o n d r i o n ( 1 1 ) .
i t w a s f o u n d t h a t t h e s i g n a l - t o - n o i s e r a t i o w a s m u c h b e t t e r f o r
t h e n e u r o n s p r o j e c t i n g u p t h e d o r s o - s p i n o - c e r e b e l l a r t r a c t( D S C T ) t o t h e c e r e b e l l u m .T h i s s u c c e s s f u l q u a n t a l r e s o l u t i o n f o r DSCT n e u r o n s a n d
m o t o n e u r o n s g i v e s c o n f i d e n c e i n t h e m u c h m o r e d i f f i c u l ta n a l y s i s o f n e u r o n s o f t h e c e r e b r a l c o r t e x , w h i c h p r o v i d e t h ek e y s t r u c t u r e s o f n e u r a l e v e n t s t h a t c o u l d b e i n f l u e n c e d b ym e n t a l e v e n t s . T h e s i g n a l - t o - n o i s e r a t i o w a s s o l o w i n t h es t u d i e s o f CA1 n e u r o n s o f t h e h i p p o c a m p u s t h a t s o f a r o n l yt h r e e q u a n t a l a n a l y s e s h a v e b e e n r e l i a b l e i n t h e c o m p l e xd e c o n v o l u t i o n p r o c e d u r e .
I n t h e m o s t r e l i a b l e a n a l y s i s , a s i n g l e a x o n o f a CA3h i p p o c a m p a l p y r a m i d a l c e l l s e t u p a n EPSP o f q u a n t a l s i z e2 7 8 A V ( m e a n v a l u e ) i n a s i n g l e CA1 h i p p o c a m p a l p y r a m i d a lc e l l w i t h a p p r o x i m a t e l y e q u a l p r o b a b i l i t i e s o f r e l e a s e a t e a c ha c t i v e s i t e ( n = 5 ) o f 0 . 2 7 ( 2 2 ) . I n t h e a l t e r n a t i v e p r o c e d u r et h e s i n g l e CA3 i m p u l s e p r o j e c t i n g t o a CA1 p y r a m i d a l c e l lw a s d i r e c t l y s t i m u l a t e d i n t h e s t r a t u m r a d i a t u m . T h e E P S P sd e l i v e r e d b y t h e d e c o n v o l u t i o n a n a l y s i s o f t w o CA1 p y r a m i -d a l c e l l s w e r e o f q u a n t a l s i z e s 2 2 4 A V a n d 1 9 3 , u V w i t hp r o b a b i l i t i e s ( n = 3 ) o f 0 . 2 4 a n d ( n = 6 ) o f 0 . 1 6 , r e s p e c t i v e l y( 2 3 ) . F o r a s y s t e m a t i c r e v i e w , s e e r e f . 2 4 .
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I n t h e w h o l e e l e c t r o p h y s i o l o g i c a l p r o c e s s o f b u i l d i n g u p t h esummed EPSP e x p e r i e n c e d i n t h e s o m a , t h e r e i s o n l y o n ee l e m e n t w h e r e q u a n t a l p r o c e s s e s c a n p l a y a r o l e . W hen ab o u t o n i s a c t i v a t e d b y a n e r v e i m p u l s e , e x o c y t o s i s o c c u r s
o n l y w i t h a c e r t a i n p r o b a b i l i t y , w h i c h i s m u c h l e s s t h a n 1 .T h i s c a l l s f o r , i n p r i n c i p l e , t h e i n t r o d u c t i o n o f t h e r m o d y -
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F I G . 2 . ( A ) D r a w i n g o f a l a m i n a V p y r a m i d a l c e l l w i t h i t s a p i c a ld e n d r i t e s h o w i n g t h e s i d e b r a n c h e s a n d t h e t e r m i n a l t u f t , a l l s t u d d e dw i t h s p i n e s y n a p s e s ( n o t a l l s h o w n ) . T h e s o m a w i t h i t s b a s a ld e n d r i t e s h a s a n a x o n w i t h a x o n c o l l a t e r a l b e f o r e l e a v i n g t h e c o r t e x .( B ) D r a w i n g o f t h e s i x l a m i n a e o f t h e c e r e b r a l c o r t e x w i t h t h e a p i c a ld e n d r i t e s o f p y r a m i d a l c e l l s o f l a m i n a e I I , I I I a n d V , s h o w i n g t h em a n n e r i n w h i c h t h e y b u n c h i n a s c e n d i n g t o l a m i n a I , w h e r e t h e y e n di n t u f t s ( A . P e t e r s , p e r s o n a l c o m m u n i c a t i o n .
n a m i c o r q u a n t u m m e c h a n i c a l s t a t i s t i c a l c o n c e p t s . W e w i s ht o m a k e i t c l e a r t h a t w e a d h e r e t o t h e q u a n t a l s t a n d p o i n t . A sh a s b e e n p o i n t e d o u t ( 1 , 2 , 5 ) , t h e c o n s c i o u s a c t i o n o f t h eb r a i n c o u l d h a r d l y b e u n d e r s t o o d i f t h e b r a i n w e r e i n i t se n t i r e t y f u n c t i o n i n g o n t h e b a s i s o f c l a s s i c a l p h y s i c s . A l s o ,t h e r e l a t i v e c o n s t a n c y o f t h e e m i s s i o n p r o b a b i l i t y f o r s i n g l eb o u t o n r e l e a s e ( 2 1 - 2 4 ) c a n h a r d l y b e e x p l a i n e d o n t h e b a s i so f t h e r m a l f l u c t u a t i o n s .
B e c a u s e t h e r e s u l t i n g EPSP i s t h e i n d e p e n d e n t s t a t i s t i c a lsum o f s e v e r a l t h o us a n ds o f l o c a l E P S P s a t s p i n e s y n a p s e s o n
e a c h d e n d r i t e ( F i g . 2 A ) , w e c a n c o n c e n t r a t e o n t h e p r o c e s so f e x o c y t o s i s a t e a c h i n d i v i d u a l b o u t o n .
E x o c y t o s i s i s t h e o p e n i n g o f a c h a n n e l i n t h e PVG a n d
d i s c h a r g e o f t h e v e s i c l e s t r a n s m i t t e r m o l e c u l e s i n t o t h e
s y n a p t i c c l e f t ( F i g . 3 F a n d G ) . I t i s , a s a w h o l e , c e r t a i n l y ac l a s s i c a l m e m b r a n e - m e c h a n i c a l p r o c e s s . To i n v e s t i g a t e f u r -t h e r t h e p o s s i b l e r o l e o f q u a n t u m m e c h a n i c s i n t h e p r o b a b i -l i s t i c d i s c h a r g e , o n e h a s t o s e t u p a m o d e l f o r t h e t r i g g e rm e c h a n i s m b y w h i c h C a 2 + p r e p a r e s t h e v e s i c l e s o f PVG f o re x o c y t o s i s .
F o r t h i s p u r p o s e w e a d o p t t h e f o l l o w i n g c o n c e p t : p r e p a -r a t i o n f o r e x o c y t o s i s m e a n s b r i n g i n g t h e p a r a c r y s t a l l i n e PVGi n t o a m e t a s t a b l e s t a t e f r o m w h i c h e x o c y t o s i s c a n o c c u r . T h et r i g g e r m e c h a n i s m i s t h e n m o d e l e d b y t h e m o t i o n o f a
q u a s i p a r t i c l e w i t h 1 d e g r e e o f f r e e d o m a l o n g a c o l l e c t i v ec o o r d i n a t e , a n d o v er a n a c t i v a t i o n b a r r i e r ( F i g . 4 ) . T h i s
m o t i o n s e t s i n b y a q u a n t u m m e c h a n i c a l t u n n e l i n g p r o c e s st h r o u g h t h e b a r r i e r ( s i m i l a r t o r a d i o a c t i v e d e c a y ) .
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F I G . 3 . ( A ) A x o n t e r m i n a l , o r b o u t o n , s h o w i n g d e n s e p r o j e c t i o n s( d p ) p r o j e c t i n g f r o m t h e a c t i v e s i t e w i t h c r o s s l i n k a g e s f o r m i n g t h eP V G , w h i c h i s d r a w n i n t h e I n s e t w i t h d i m e n s i o n s ( 1 4 ) . c v , s m a l lv e s i c l e ; d v , b i g v e s i c l e . ( B ) T a n g e n t i a l s e c t i o n t h r o u g h t h e p r e s y n -a p t i c a r e a . A s e c t i o n o f t h e p a t t e r n s o f d e n s e p r o j e c t i o n s a n d s y n a p t i cv e s i c l e s , t r i a n g u l a r a n d h e x a g o n a l , o f t h e PV G i s c l e a r l y r e p r e s e n t e d .( B a r = 0 . 1 g m . ) [ R e p r i n t e d f r o m r e f . 1 5 w i t h p e r m i s s i o n . ] ( C - E )A c t i v e z o n e ( A Z ) o f m a m m a l i a n c e n t r a l s y n a p s e s h o w i n g g e o me t -r i c a l d e s i g n ( 1 1 ) . S V , s y n a p t i c v e s i c l e ; V A S , v e s i c l e a t t a c h m e n t s i t e ;P A , p r e s y n a p t i c a r e a . ( F ) S y n a p t i c v e s i c l e i n a p p o s i t i o n . ( G ) E x o -c y t o s i s ( 1 6 ) .
T h e s i t u a t i o n i s d e t e r m i n e d b y t w o c h a r a c t e r i s t i c e n e r g i e s : i ) t h e t h e r m a l e n e r g y , E t h , w h i c h t h e q u a s i p a r t i c l e a d o p t s i na t h e r m a l s u r r o u n d i n g o f t e m p e r a t u r e T . I t i s f o r o u r q u a s i p a r -t i c l e w i t h 1 d e g r e e o f f r e e d o m g i v e n b y
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l o c a l i z e d o v e r a d i s t a n c e o f A q . I t f o l l o w s f r o m H e i s e n b e r g s
u n c e r t a i n t y r e l a t i o n A p - A q z 2 1 r h . I f w e a d o p t t h e l o w e r l i m i t( t h i s w o u l d c o r r e s p o n d t o t h e g r o u n d s t a t e i n a c o n f i n i n gp o t e n t i a l w e l l ) , w e o b t a i n
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F I G . 4 . T h e c o l l e c t i v e p o t e n t i a l , V ( q ) , f o r t h e m o t i o n o f t h eq u a s i p a r t i c l e o f e n e r g y E o w h i c h t r i g g e r s e x o c y t o s i s . T h e d a s h e dc u r v e s k e t c h e s a t u n n e l i n g s t a t e t h r o u g h t h e b a r r i e r . A t t h e b e g i n n i n gt h e w a v e p a c k e t i s l o c a t e d i n t h e l e f t a r e a . A f t e r t i m e T t h e a m p l i t u d e
h a s a p a r t l e f t o f t h e b a r r i e r a n d a p a r t r i g h t o f t h e b a r r i e r .
W e c a n d e f i n e a b o r d e r l i n e b e t w e e n t h e q u a n t a l a n d t h e
c l a s s i c a l r e g i m e b y p u t t i n g E o = E t h . E o >> E t h i s t h e n t h eq u a n t a l , a n d E c h >> E o t h e t h e r m a l r e g i m e .
F o r f i x e d T a n d Aq t h e b o r d e r l i n e e q u a l i t y d e t e r m i n e s a
c r i t i c a l m a s s , M C , f o r t h e q u a s i p a r t i c l e . D y n a m i c p r o c e s s e sw h i c h i n v o l v e q u a s i p a r t i c l e m a s s e s much l a r g e r t h a n MCb e l o n g t o t h e c l a s s i c a l r e g i m e , w h e r e a s f o r M << MC w e a r e
i n t h e q u a n t a l r e g i m e . T a k i n g T = 3 0 0 K a n d Aq = 1 A , w eo b t a i n MC - 1 0 - 2 3 g = 6 MH, w i t h MH b e i n g t h e m a s s o f a
h y d r o g e n a t o m .
T h i s e s t i m a t e s h o w s q u i t e c l e a r l y t h a t a q u a n t u m m ec h a n -i c a l t r i g g e r f o r e x o c y t o s i s m u s t r e s i d e i n a n a t o m i c p r o c e s s -e . g . , t h e m o v e m e n t o f a h y d r o g e n b r i d g e b y e l e c t r o n i cr e a r r a n g e m e n t .
To m a k e t h e m o d e l q u a n t i t a t i v e , w e a t t r i b u t e t o t h e t r i g -g e r i n g p r o c e s s o f e x o c y t o s i s a c o n t i n u o u s c o l l e c t i v e v a r i a b l e ,q , f o r t h e q u a s i p a r t i c l e . T h e m o t i o n i s c h a r a c t e r i z e d b y a
p o t e n t i a l e n e r g y , V ( q ) , w h i c h may t a k e o n a p o s i t i v e v a l u e a ts t a g e I ( F i g . 4 ) , a c c o r d i n g t o t h e m e t a s t a b l e s i t u a t i o n b e f o r e
e x o c y t o s i s ,t h e n r i s e s t o w a r d a maximum
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f i n a l l y d r o p s t o z e r o ( t h e a r b i t r a r y n o r m a l i z a t i o n ) a t s t a g e I V .F i g . 4 g i v e s a q u a l i t a t i v e s ke t c h o f t h i s p o t e n t i a l b a r r i e r .
T h e t i m e - d e p e n d e n t p r o c e s s o f e x o c y t o s i s i s d e s c r i b e d b yt h e o n e - d i m e n s i o n a l S c h r o d i n g e r e q u a t i o n f o r t h e wave f u n c -t i o n q ( q ; t ) ,
a / i 2 a 2i h q , q ; t ) O f q ; t ) V ( q ) * ( q ; t ) . [ 3 ]
a t 2Ma q 2
T h e i n i t i a l c o n d i t i o n f o r t = 0 ( s t a g e I , b e g i n n i n g o f e x o c y -t o s i s ) i s a w a v e p a c k e t t o t h e l e f t o f t h e p o t e n t i a l b a r r i e r ( F i g .4 ) .
T h e s o l u t i o n o f E q . 3 g i v e s t h e wave f u n c t i o n a f t e r t i m e t ,w h i c h c o n s i s t s o f p a r t o f t h e w a v e p a c k e t s t i l l r e s i d i n g t o t h el e f t o f t h e b a r r i e r w h i l e a n o t h e r p a r t h a s p e n e t r a t e d i n t o t h es p a c e t o t h e r i g h t ( s e e F i g . 4 ) . T h e q u a n t i t i e s
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o c c u r ( P l ) o r t h a t e x o c y t o s i s w i l l o c c u r ( b y a c t i o n o f t h et r i g g e r ) P 2 ) . O f c o u r s e , f r o m n o r m a l i z a t i o n , p i ( t ) + p 2 ( t ) =1 .
T h e r e l e v a n t t i m e r h a s t o b e d e f i n e d a s t h e d u r a t i o n o f t h eq u a s i s t a b l e s i t u a t i o n i n t h e p a r a c r i s t a l l i n e P V G , a f t e r w h i c h
t h e m e t a s t a b l e s t a t e c h a n g e s i n t o a s t a b l e o n e w h e r e n o
e x o c y t o s i s i s p o s s i b l e . I n o u r m o d e l t h i s c a n b e v i s u a l i z e d a s
a d r o p p i n g o f t h e p o t e n t i a l l e f t o f t h e b a r r i e r t o a v a l u e f o rw h i c h b a r r i e r p e n e t r a t i o n h a s v a n i s h i n g l y s m a l l p r o b a b i l i t y .An a p p r o x i m a t e s o l u t i o n o f t h e b a r r i e r p e n e t r a t i o n p r o b -
l e m c a n b e o b t a i n e d f r o m t h e W e n t z e l - K r a m e r s - B r i l l o u i n WKB) m e t h o d ( 2 5 ) . A c c o r d i n g t o i t t h e t r a n s m i s s i o n c o e f -f i c i e n t T o f a p a r t i c l e o f m a s s M a n d e n e r g y E t h r o u g h t h eb a r r i e r i s g i v e n b y
T = exP{-2f 2 M [ V ( q ) E ] [ 5 ]
w h e r e a a n d b d en o t e t h e c l a s s i c a l t u r n i n g p o i n t s f o r t h e
m o t i o n l e f t a n d r i g h t o f t h e b a r r i e r ( F i g . 4 ) . T h e p r o b a b i l i t y
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p e r u n i t t i m e f o r p e n e t r a t i o n o f t h e b a r r i e r , w , c a n b e o b t a i n e db y t h e n u m b e r o f a t t e m p t s t h a t t h e p a r t i c l e u n d e r t a k e s t or e a c h t h e b a r r i e r , c , , t i m e s t h e b a r r i e r t r a n s m i s s i o n c o e f f i -c i e n t , T ,
d p o T . 1 ~ ~ ~ ~ ~ 6 ]d t
w i t h h / i o = E O b y a n a p p e a l t o B o h r s c o r r e s p o n d e n c ep r i n c i p l e . C o m b i n i n g t h i s w i t h t h e g e n e r a l c o n s i d e r a t i o n sf o r m u l a t e d a f t e r E q . 4 , we o b t a i n
P 2 ( T ) = x*@= T w o W T . [ 7 ]
A t t h i s s t a g e we c a n e s t i m a t e s o m e n u m b e r s . L et u sa s s u m e t h a t t h e e n e r g y E = E o o f t h e i n i t i a l s t a t e ( t h e w a v ep a c k e t l o c a l i z e d t o t h e l e f t o f t h e b a r r i e r ) i s t h e z e r o - p o i n te n e r g y o f a w a v e p a c k e t l o c a l i z e d o v e r t h e d i m e n s i o n o f t h ea t o m i c s i t e o f t h e t r i g g e r , w h i c h w e e s t i m a t e r o u g h l y t o b e A q
1 A , a n d l e t u s t a k e f o r t h e m a s s M o f o u r q u a s i p a r t i c l e t h em a s s o f o n e h y d r o g en a t o m , M = 1 . 7 x 1 0 - 2 4 g . T h e n w eo b t a i n , u s i n g E q . 2 , E o = 8 . 3 x 1 0 - 2 e V . T h i s l e a d s t o a
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1 . 3 x 1 0 1 4 s - 1 . E s t i m a t i n gx , t h e t i m e o f t h e m e t a s t a b l e i n s t a b i l i t y , t o b e o f t h e o r d e r o fe l e c t r o n i c t r a n s i t i o n p r o c e s s e s w h i c h i s r 1 0 - 1 3 - 1 0 - 1 4 s w eo b t a i n f r o m E q . 7 P 2 ( T ) 10 o T t o 1 0 0 O T . W i t h t h e o b s e r v e d P 2o f a b o u t 0 . 2 5 , t h i s g i v e s f o r t h e b a r r i e r p e n e t r a t i o n f a c t o r Tt h e r e a s o n a b l e s p a n o f 4 x 1 0 - 2 t o 4 x 1 0 - 3 .Up t o n o w w e h a v e b a s e d o u r m o d e l c o n s i d e r a t i o n s o n t h e
r e l e a s e o f o n e s i n g l e v e s i c l e l o c a t e d i n t h e v e s i c u l a r g r i d o f t h eb o u t o n . T h e r e a r e a b o u t 4 0 v e s i c l e s a l t o g e t h e r i n t h i sp a r a c r y s t a l l i n e s t r u c t u r e , b u t n ev e r d o e s m o r e t h a n o n ev e s i c l e e m i t t r a n s m i t t e r m o l e c u l e s i n t o t h e s y n a p t i c c l e f t a f t e rs t i m u l a t i o n b y a n er v e i m p u l s e . T h i s c e r t a i n l y m e a n s t h a t t h ev e s i c l e s i n t h e v e s i c u l a r g r i d d o n o t a c t i n d e p e n d e n t l y , b u tr a t h e r t h a t i m m e d i a t e l y a f t e r o n e v e s i c l e i s t r i g g e r e d f o rr e l e a s i n g i t s c o n t e n t t h e i n t e r a c t i o n b e t w e e n t h e m b l o c k sf u r t h e r e x o c y t o s i s . T h e p a r a c r y s t a l l i n e s t r u c t u r e o f t h e PVGm a k e s i t p o s s i b l e t o h a v e l o n g - r a n g e i n t e r a c t i o n s b e t w e e n t h ec o n s t i t u e n t s , a s i s w e l l k n o w n f r o m o r d e r e d q u a n t u m s y s -t e m s . A c c o r d i n g t o o u r n u m e r i c a l e s t i m a t e s , t h e r e l a x a t i o nt i m e f o r t h e b l o c k i n g p r o c e s s i s o f t h l e o r d e r o f f e m t o s e c o n d s .
W i t h t h i s o b s e r v a t i o n we c a n d i s c u s s t h e m a n y - b o d ya s p e c t o f e x o c y t o s i s f r o m t h e v e s i c u l a r g r i d . T o t h i s e n d w ea t t r i b u t e s c h e m a t i c a l l y t o e a c h v e s i c l e i n t h e g r i d t w o s t a t e s ,
4 f a n d 4 i , w h e r e i i s t h e s t a t e b e f o r e a n d I i t h e s t a t e a f t e re x o c y t o s i s h a s b e e n t r i g g e r e d . W e c a n s a f e l y a s s u m e t h a t t h ed i f f e r e n t v e s i c l e s a r e s o w e l l s e p a r a t e d t h a t w e c a n t r e a t t h e ma s d i s t i n g u i s h a b l e p a r t i c l e s . T h e w a v e f u n c t i o n o f N v e s i c l e si s t h e n a p r o d u c t o f d e n u m e r a b l e s t a t e s ,
t V ,N)= ( 2 ) ( N ) { o , 1 } . [ 8
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o ( 1 , . . . N ) = o l ) q 4 N ) . [ 9 ]
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A l l o t h e r s t a t e s a r e p u s h e d u p i n e n e r g y b y t h e l o n g - r a n g ei n t e r a c t i o n s o f a r t h a t t h e y c a n n o t b e e x c i t e d b y t h e n e r v ei m p u l s e .
C a l c u l a t i n g t h e p r o b a b i l i t y f o r e x o c y t o s i s f r o m t h e N - b o d yw a v e f u n c t i o n s , E q s . 9 a n d 1 0 , o n e o b t a i n s t h e s a m e r e s u l t a sw a s o b t a i n e d f r o m t h e b a r r i e r - p e n e t r a t i o n p r o b l e m o f o n ev e s i c l e , s i n c e t h e t r i g g e r c a n c h a n g e t h e s t a t e o f o n l y o n ev e s i c l e , b u t t h e r e a r e N s u c h p o s s i b i l i t i e s . T h i s l e a d s t o t h e
o b s e r v a b l e c o n s e q u e n c e t h a t t h e p r o b a b i l i t y f o r e x o c y t o s i s o fo n e b o u t o n d o e s n o t d e p e n d o n t h e n u m b e r o f i d e n t i c a lv e s i c l e s o c c u p y i n g t h e P V G .
O u r m o d e l i n t r o d u c e s i n t o t h e a c t i v i t y o f t h e n eo c o r te x aq u a n t u m p r o b a b i l i s t i c a s p e c t t h a t l e a d s t o a s e l e c t i o n o fc h o i c e s a c c o r d i n g t o a q u a n t u m p r o b a b i l i t y a m p l i t u d e .
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I n g v a r ( 2 6 ) i n t r o d u c e d t h e t er m p u r e i d e a t i o n , w h i c h i sd e f i n e d a s c o g n i t i v e e v e n t s t h a t a r e u n r e l a t e d t o a n y o n g o i n gs e n s o r y s t i m u l a t i o n o r m o t o r p e r f o r m a n c e s . H e a n d a s s o c i -a t e s a t t h e U n i v e r s i t y o f L u n d i n t r o d u c e d t h e s t u d y o f t h er e g i o n a l c e r e b r a l b l o o d f l o w t o d i s p l a y b y c e r e b r a l i d e o g r a -p h y t h e a c t i v i t y o f t h e b r a i n i n p u r e i d e a t i o n i n a l l t h e
i m m e n s e v a r i e t y g e n e r a t e d b y t h e p s y c h e . By r a d i o x e n o nm a p p i n g R o l a n d e t a l . ( 2 7 ) d e m o n s t r a t e d t h a t i n p u r e m o t o ri d e a t i o n o f c o m p l e x h a n d m o v e m e n t s t h e r e w a s a c t i v a t i o n o ft h e s u p p l e m e n t a r y m o t o r a r e a ( S M A ) o n b o t h s i d e s . B y t h em o r e a c c u r a t e t e c h n i q u e o f p o s i t r o n e m i s s i o n t o m o g r a p h y( P E T ) s c a n n i n g R a i c h l e a n d h i s a s s o c i a t e s ( 2 8 ) d e m o n s t r a t e da w i d e s p r e a d p a t c h y a c t i v i t y o f t h e n e o c o r t e x d u r i n g s p e c i f i cm e n t a l o p e r a t i o n s i n s e l e c t i v e a t t e n t i o n .
I n a c o m p l e m e n t a r y p r o c e d u r e , a v e r a g i n g t e c h n i q u e s h a v eb e e n u s e d t o r e c o r d t h e e l e c t r i c a l f i e l d s g e n e r a t e d b y t h eb r a i n ( c e n t e r e d o n t h e SMA) i n t h e w i l l i n g o f a m o v e m e n t , t h es o - c a l l e d r e a d i n e s s p o t e n t i a l ( 2 9 ) . I n e x q u i s i t e l y d e s i g n e de x p e r i m e n t s L i b e t ( 3 0 ) h a s d i s c o v e r e d t h a t i n c o n s c i o u sw i l l i n g t h e r e i s a c e r e b r a l a c t i v a t i o n a bo u t 2 0 0 m s b e f o r e t h em o v e m e n t ( s e e r e f . 3 0 , G e n e r a l D i s c u s s i o n ) .
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c o r t i c a l n e u r o n , a h i p p o c a m -p a l p y r a m i d a l c e l l , d i s c l o s e s a c o n t i n u o u s i n t e n s e a c t i v i t yw h i c h c a n b e i n t e r p r e t e d a s m i l l i - E P S P s g e n e r a t e d b y t h ec o n t i n u o u s s y n a p t i c b o m b a r d m e n t w i t h e x o c y t o s e s b y t h et h o u s a n d s o f b o u t o n s o n i t s d e n d r i t e s ( F i g . 2 A ) . D e c o n v o -l u t i o n a n a l y s i s h a s s h o w n t h a t a n i m p u l s e i n v a d i n g a b o u t o n( F i g . 3 A a n d D ) e v o k e s a n e x o c y t o s i s a n d a m i l l i - E P S P w i t ha p r o b a b i l i t y o f a b o u t 0 . 2 - 0 . 3 ( 2 2 , 2 3 ) .
C o m b i n i n g t h e s e o b s e r v a t i o n s w i t h o u r q u a n t u m m e c h a n -i c a l a n a l y s i s o f b o u t o n e x o c y t o s i s , w e p r e s e n t now t h eh y p o t h e s i s t h a t t h e m e n t a l i n t e n t i o n ( t h e v o l i t i o n ) b e c o m e sn e u r a l l y e f f e c t i v e b y m o m e n t a r i l y i n c r e a s i n g t h e p r o b a b i l i t yo f e x o c y t o s i s i n s e l e c t e d c o r t i c a l a r e a s s u c h a s t h e S M An e u r o n s ( 3 1 ) . I n t h e l a n g u a g e o f q u a n t u m m e c h a n i c s t h i sm e a n s a s e l e c t i o n o f e v e n t s ( t h e e v e n t t h a t t h e t r i g g e rm e c h a n i s m h a s f u n c t i o n e d , w h i c h i s a l r e a d y p r e p a r e d w i t h a
c e r t a i n p r o b a b i l i t y ; c f . F i g . 4 a n d E q . 4 ) . T h i s a c t o f s e l e c t i o ni s r e l a t e d t o W i g n e r s s e l e c t i o n p r o c e s s o f t h e m i n d o nq u a n t a l s t a t e s 6 ) , a n d i t s m e c h a n i s m c l e a r l y l i e s b e y o n do r d i n a r y q u a n t u m m e c h a n i c s . E f f e c t i v e l y t h i s s e l e c t i o nm e c h a n i s m i n c r e a s e s t h e p r o b a b i l i t y f o r e x o c y t o s i s , a n d i nt h i s w a y g e n e r a t e s i n c r e a s e d E P S P s w i t h o u t v i o l a t i o n o f t h ec o n s e r v a t i o n l a w s . F u r t h e r m o r e , t h e i n t e r a c t i o n o f m e n t a le v e n t s w i t h t h e q u a n t u m p r o b a b i l i t y a m p l i t u d e s f o r e x o c y -t o s i s i n t r o d u c e s a c o h e r e n t c o u p l i n g o f a l a r g e n u m b e r o fi n d i v i d u a l a m p l i t u d e s o f t h e h u n d r e d s o f t h o u s a n d s o f b o u -t o n s i n a d e n d r o n . T h i s t h e n l e a d s t o a n o v e r w h e l m i n g v a r i e t yo f a c t u a l i t i e s , o r m o d e s , i n b r a i n a c t i v i t y . P h y s i c i s t s w i l lr e a l i z e t h e c l o s e a n a l o g y t o l a s e r - a c t i o n , o r , m o r e g e n e r a l l y ,t o t h e p h e n o m e n o n o f s e l f - o r g a n i z a t i o n .
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e x p l a i n e d b y n e u r o s c i e n c e i n t h e c o n v e n t i o n a l m a n n e r .
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T h e r e c o ul d b e s u m m a t i o n o f t h e m i l l i - E P S P s g e n e r a t e d b yt h e t h o u s a n d s o f b o u t o n s o n e a c h p y r a m i d a l c e l l ( F i g . 2 A ) ,
a n d t h e n t h e h u n d r e d o r s o p y r a m i d a l c e l l s o f a d e n d r o n ( F i g .2 B ) w o u l d i n t e r a c t i n g e n e r a t i n g t h e o n g o i n g d i s c h a r g e o f t h eS M A n e u r o n s ( 3 1 ) , a n d f u r t h e r t o t h e m o t o r p y r a m i d a l c e l l sw i t h d i s c h a r g e s down t h e p y r a m i d a l t r a c t t o t h e m o t o n e u -r o n s . T h e r e s u l t i n g i m p u l s e d i s c h a r g e s i n m o t o r n e r v e s w o u l de f f e c t t h e i n t e n d e d m o v e m e n t w i t h a l l t h e c o n t r o l s y s t e m s
c o o p e r a t i n g .V o l u n t a r y m o v e m e n t h a s b e e n e x p l a i n e d i n p r i n c i p l e . T h i s
e x p l a n a t i o n c a n b e e x t e n de d t o t h e a c t i o n o f a l l m e n t a li n f l u e n c e s o n t h e b ra i n - f o r e x a m p l e, i n c a r r y i n g o u t a n yp l a n n e d a c t i o n , s u c h a s s p e e c h .
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B a s e d o n a c a r e f u l a n a l y s i s o f n e o c o r t i c a l a c t i v i t y , we a r g u et h a t e x o c y t o s i s i s i t s k e y m ec h a n i s m. E x o c y t o s i s , t h e m o -m e n t a r y o p en i n g o f a c h a n n el i n t h e p r e s y n a p t i c m e m b r a n eo f a b o u t o n w i t h l i b e r a t i o n o f t h e t r a n s m i t t e r s u b s t a n c e ( F i g .3 F a n d G ) , i s c a u s e d b y a n e r v e i m p u l s e . A s h a s b e e ne s t a b l i s h e d i n m a n y e x p e r i m e n t s ( 2 2 - 2 4 ) , e x o c y t o s i s i s a na l l - o r - n o t h i n g - e v e n t , o c c u r r i n g w i t h p r o b a b i l i t i e s o f t h e o r d e r
o f 0 . 2 5 . T h i s o b s e r v a t i o n l e d u s t o s e t u p a q u a n t u m m ec h a n -i c a l m o d e l f o r t h e t r i g g e r m e c h a n i s m o f e x o c y t o s i s , b a s e d o nt h e t u n n e l i n g o f a q u a s i p a r t i c l e r e p r e s e n t i n g t h e t r i g g e r .
T h e q u a n t u m t r e a t m e n t o f e x o c y t o s i s l i n k s t h e n e o c o r t i c a la c t i v i t y w i t h t h e e x i s t e n c e o f a l a r g e n u m b e r o f q u a n t u mp r o b a b i l i t y a m p l i t u d e s , s i n c e t h e r e a r e m o r e t h a n 1 0 0 , 0 0 0b o u t o n s i n a b u n d l e o f d e n d r i t e s c a l l e d a d e n d r o n ( F i g . 2 Aa n d B ) . I n t h e a b s e n c e o f m en t a l a c t i v i t y t h e s e p r o b a b i l i t ya m p l i t u d e s a c t i n d e p e n d e n t l y , c a u s i n g f l u c t u a t i n g E P S P s i nt h e p y r a m i d a l c e l l . W e p u t f o r w a r d t h e h y p o t h e s i s t h a tm e n t a l i n t e n t i o n b e c o m e s n e u r a l l y e f f e c t i v e b y m o m e n t a r i l yi n c r e a s i n g t h e p r o b a b i l i t i e s f o r e x o c y t o s e s i n a w h o l e d e n -d r o n ( F i g . 2 B ) a n d , i n t h i s w a y , c o u p l e s t h e l a r g e n u m b e r o f
p r o b a b i l i t y a m p l i t u d e s t o p r o d u c e c o h e r e n t a c t i o n .O u r h y p o t h e s i s o f f e r s a n a t u r a l e x p l a n a t i o n f o r v o l u n t a r y
m o v e m e n t s c a u s e d b y m e n t a l i n t e n t i o n s w i t h o u t v i o l a t i n gp h y s i c a l c o n s e r v a t i o n l a w s . I t h a s b e e n s h o w n e x p e r i m e n -
t a l l y t h a t i n t e n t i o n s a c t i v a t e t h e c e r e b r a l c o r t e x i n c e r t a i nw e l l - d e f i n e d r e g i o n s p r i o r t o t h e m o v e m e n t ( 2 6 - 2 8 ) .
T h e u n i t a r y h y p o t h e s i s t r a n s f o r m s t h e m a n n e r o f o p e r a t i o no f t h e i n t e n t i o n . O n e h a s t o r e c o g n i z e t h a t , i n a l i f e t i m e o fl e a r n i n g , t h e i n t e n t i o n t o c a r r y o u t a p a r t i c u l a r m o v e m e n tw o u l d b e d i r e c t e d t o t h o s e p a r t i c u l a r d en d ro n s o f t h e n e o -
c o r t e x t h a t a r e a p p r o p r i a t e f o r b r i n g i n g a b o u t t h e r e q u i r e da c t i o n s . W e b e l i e v e t h a t t h e p r o p o s e d h y p o t h e s i s a c c o u n t sf o r a c t i o n a c r o s s t h e m i n d - b r a i n i n t e r f a c e .
W e a c k n o w l e d g e t h e h e l p o f D r . S t e p h e n Redman i n c a l c u l a t i o n so f p r o b a b i l i t i e s o f r e l e a s e .
1 . S t a p p , H . P . ( 1 9 9 1 ) F o u n d . P h y s i c s 2 1 , 1 4 5 1 - 1 4 7 7 .2 . S q u i r e s , E . J . ( 1 9 8 8 ) F o u n d . P h y s i c s L e t t . 1 , 1 3 - 2 0 .3 . D o n a l d , M . J . ( 1 9 9 0 ) P r o c . R . S o c . L o n d o n A 4 2 4 , 4 3 - 9 3 .4 . M a r g e n a u , H . ( 1 9 8 4 ) T h e M i r a c l e o f E x i s t e n c e ( O x B o w ,
W o o d b r i d g e , C T ) .
5 . E c c l e s , J . C . ( 1 9 9 0 ) P r o c . R . S o c . L o n d o n B 2 4 0 , 4 3 3 - 4 5 1 .
6 . W i g n e r , E . P . ( 1 9 6 7 ) i n S y m m e t r i e s a n d R e f l e c t i o n s ( I n d i a n aU n i v . P r e s s , B l o o m i n g t o n , I N ) , p p . 1 5 3 - 1 8 4 .
7 . v o n N e u m a n n , J . ( 1 9 5 5 ) M a t h e m a t i c a l F o u n d a t i o n s o f Q u a n -
t u m M e c h a n i c s ( P r i n c e t o n U n i v . P r e s s , P r i n c e t o n ) .8 . E v e r e t t , H . ( 1 9 5 7 ) R e v . M o d . P h y s . 2 9 , 4 5 4 - 4 6 2 .
9 . P o p p e r , K . E c c l e s , J . C . ( 1 9 7 7 ) T h e S e l f a n d I t s B r a i n
( S p r i n g e r , B e r l i n ) .1 0 . S z e n t a g o t h a i , J . ( 1 9 7 8 ) P r o c . R . S o c . L o n d o n B 2 0 1 , 2 1 9 - 2 4 8 .
1 1 . G r a y , E . G . ( 1 9 8 2 ) T r e n d s N e u r o s c i . 5 , 5 - 6 .
1 2 . P e t e r s , A . K a r a , D . A . ( 1 9 8 7 ) J . C o m p . N e u r o l . 2 6 0 , 5 7 3 - 5 9 0 .
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B i o p h y s i c s : B e c k a n d E c c l e s
