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BpC`DEF166hhhhpppppph h h(h(h0h0h8h8h@h@`@`@`H`HXPXPXXXXPXPXHXHX@X@X@`@`8`8`0`0`(`(` ` `````````XXmp@" @ BpC`DEF1iihhhhpppppppppppppph h h h h h(h(h(h(h(h(h0h0h0h0h0h8h8h8h8h8h@`@`@`H`H`H`H`HXPXPXPXPXXPXPXPXPXPXPXPXHXHXHX@X@X@X@X@X@`8`8`8`8`8`0`0`0`(`(`(` ` ` ` ` ```````````````````XXX@"& A BC0DEF100  ( (((((0(0(00008080@0@0H0H0P0P0X0X0X(X(`(`(h(h(p(p(p p x x ad@" v B 8BC0DEhFp1ZZ  (((((((((0(0(00808080@0@0@0@0@0H0H0H0H0H0P0P0P0P0P0P0X0X0X0X(X(X(`(`(`(`(`(h(h(h(h(h(h(h(p(p p p p x x x x @" 6 C BC(DEHFP1(( %(@"H@Xh D `BC(DE|F1(((( ?@@"H@Xh6 E BC DEHFP1      %(@"Hh F BC DEF1$$       IL@"Hh2 G xE$GHI`TQ1 `T`T`T`T`T`T`T`T`T`T" 2 H xE$GHI`TQ1 `T`T`T`T`T`T`T`T`T`T"0V2 I C G1" V2 J C G1"p  K BCDEF 1@@`` @"( L BCDE F1@`@"( M BCDEF 1@@hh @"   N BCDE F1@h@"  ~T 4   O# H  P  ~BxC`DEF1!!xxhXPH@0( 08@@HXX`` X(X(X0P@HH@X8h(p p xxxxDH@`"4  V2 Q C 1"L  tT \  R#  S B8CpDEF p 8 p @`"\ JB T # 1"| } n U 0A#?" wSn V 0A'?"M9T g  W# As  X zBCDE F1@"_ V Y C 1"#g V Z C 1"g V [ C 1": pg V \ C 1"h g V ] C 1" Ig  _ Z\ ?? p F" Development: The formation and refinement of neural circuits need synaptic elimination. Axonal or synaptic competition in neuromuscular junctions and visual system (Consumptive and interference competition) L@S@u@ @`H  0޽h ? "-. / 0 G H IJ ̙3380___PPT10.5f 0 (  T  Z ??P0 " Long term potentiation (LTP) - Long term depression LTD Changes that persist for tens of minutes or longer are generally called LTP and LTD. It lasts for hours in vitro and days and weeks in vivo The longest-lasting forms appear to require protein synthesis. First found in Hippocampus The physiological basis of Hebbian learning Properties and mechanisms of long-term synaptic plasticity in the mammalian brain may relate to learning and memory. Inhibitory synapses can also display plasticity, but this has been less thoroughly investigated both experimentally and theoreticallya5@`       @  @ N   NI @`H  0޽h ? ̙3380___PPT10.òЃY" 0  "(    6 p>6.___PPT9 >Introduction Synaptic placticity rules " The basic Hebb rule " The covariance rule " BCM Rule " Non-Hebbian rules " Anti-hebbian rules " Timing-based Rules   Ggc c cccc c $$$$ $$(g(,G,0g0 4g48g8??  H0 ZA ?1 ?0 ~B h $0c0 # K Z\i ?? D<4___PPT9 MThe processes of synaptic plasticity are typically much slower than the neural activity dynamics. If, in addition, the stimuli are presented slowly enough to allow the network to attain its steady-state activity during training,6 ! @` M Z f??`H  0޽h ? ̙33___PPT10.>"+7uDb'  = @B D' = @BA?%,( < +O%,( < +DT' =%(%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*%(+4 0  44(  4 4 Zj ??P  hlTheoretically, an upper saturation constraint must be imposed to avoid unbounded growth. But experimentally,mcc  ccP    X 4 0A?pL 4 Z??C"?p0 PH 4 0޽h ? ̙3380___PPT10.u 0 tl -(   ) 6̊   DIs it due to that the basic Hebb rule has no LTD? Let s add LTD by introducing the covariance ruleBcO!Cv * NA ??u  , H' `   H  0޽h ? ̙3380___PPT10..  0 |t   (      6x p>6.___PPT9 (Introduction Synaptic placticity rules " The basic Hebb rule " The covariance rule " BCM Rule " Non-Hebbian rules " Anti-hebbian rules " Timing-based Rules   Ggc c cccc c $$ $$(g(,C,0c0 4g48g8??   f ??0` :P 0the input covariance matrix,pB  HD!ԔP   Z f??@ PPH  0޽h ? ̙33~v___PPT10V.`v+#^D*' = @B D' = @BA?%,( < +O%,( < +D' =%(%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* %(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*%(+ 0   LT (  LH L 0p ~v0___PPT106___PPT9 b" By the basic Hebb rule, synapses are modified whenever correlated pre- and postsynaptic activity occurs. Such correlated activity can occur purely by chance, rather than reflecting a causal relationship that should be learned. To correct for this, the covariance rather than correlation-based rule is often used by network modelshM 2eg'3gc>  3    L0 TA ? ? H $D0 T  L 0|P   " Although the covariance rule allows LTD and reflects a causal pre- and postsynaptic relationship it is still unstable due to positive feedbackt 2d( c+d`.  < )   , .  L0 TA ? ? nH $D0  L  fp ??  O(homosynaptic depression)   L Z| ??  Q(heterosynaptic depression) H L 0޽h ? ̙33___PPT10r.p1+Qb=DF' = @B D' = @BA?%,( < +O%,( < +D8' =%(%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<* L%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*L%(+ 0   &D (  Dx D c $     D0 TA ? ?0 9    D0 TA ? ?0 H $D0 O D ZB ?? p  1Average above equation over the training period: H2cc  (  r2 D s *3ffC"?P0pB D HDԔhr2 D s *3fC"?P D 0HB ?post  ` D c $A ??P7  D 0@B  ?pre  2 D Z };T?? r2 D s *3fC"?` r2  D s *3fC"?`@r2 !D s *3fC"?pB "D HDԔp8pB #D HDԔ0 pB $D HDԔ `y &D ZB ??  GCompetition can be introduced to allowing threshold to slide as followsLHc Cc< C 8    ; H D 0޽h ? ̙33___PPT10.d6+@ LDb' = @B D' = @BA?%,( < +O%,( < +DT' =%(%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*D%(+ 0 x0$(  $ $ 6*B p>6.___PPT9 ,Introduction Synaptic placticity rules " The basic Hebb rule " The covariance rule " BCM Rule " Non-Hebbian rules " Anti-hebbian rules " Timing-based Rules   Ggc c cccc c $$ $$(g(,G,0g0 4g48c8??pp `0 TA! ? ? H $D0 BR ` s *X99?  ` B1 CsDEFuj' j1 j1 s @u  ` B: CsDEFui0 i: i: s @ :B ` 3 ul l :B ` 3 ul 1 a l :B ` 3 ul l :B ` 3 ul ( X l :B ` 3 ul l :B ` 3 ul  N l :B ` 3 ul l :B  ` 3 ul El :B !` 3 ul l :B "` 3 ul ;l :B #` 3 ul l :B $` 3 ul 2l :B %` 3 ul ~l :B &` 3 ul )l :B '` 3 ul ul :B (` 3 ul l :B )` 3 ul kl :B *` 3 ul l :B +` 3 ul bl :B ,` 3 ul  l :B -` 3 ul Yl b .` :B CDElFturKqU&h/{&hLVLC{:B' 78@  :B /` 3 u  F 0` S u m  1` 6lB vp  rqyRF 2` S uF 2   3` <oB3 <   CLTP$G F 4` S u  5` <` S uF;Y ?` <B4E4 UPostsynaptic activity$G  @` ZB ?? w   One example: H g!g!!   A` 6BP 7  Usually set:8 2 c$    B D`  `D!>??   B E`  `D8Ԕ?? p B F`  `D8Ԕ??@   G` ZxB ??    Hebb rule : cc"   H` ZB ??`g Ycovariance rule.c I` Z+B#style.visibility<* `%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*`%(+  0   :It(  tn t 0B  This is again unstable if q is fixed. However, if the threshold is allowed to grow faster than v we get stability. y 2a  ` ``* l  &      * >T  -  t "s T t c $X99? -  t ~BV CDEFliN V V  @e    t ~B_ CDEFliV _ _  @n  <B  t # lie E E<B  t # li E E<B  t # liG Es E<B  t # li E E<B t # li) EU E<B t # li E E<B t # li E7E<B t # li|EE<B t # liEE<B t # li^EE<B t # liEE<B t # li@ElE<B t # liEE<B t # li"ENE<B t # liEE<B t # liE/E<B t # liuEE<B t # liEE<B t # liWEE<B t # liEE<B t # li9EdEN t BCDEdFlli>+hfJF]r) a0Gq= #Cf 34@n N <B  t # liNVH !t C li"t "t 6(Bb Fq(H #t C li# 2  $t 6$B, !  CLTP$G H %t C li= 8   &t 6BG '  CLTD$G H 't C li ;  (t 6B . } A0$G P@ (  . +t(  . )t BiCDEF75i55 @`(  .<B *t # li] ] .P@ ~q $  .t~q $  ,t BCiDEF75i55 @`q $ <B -t # li~  H /t C liP   0t 6+F]Nwhw 'Jav0q=$ 5O`-0@n ! BB 2tB 3 Nli ! !f 3t 6B CDEpFxli>Eh fJw`1TC)Oa-0)q=#.b  9<@w ! BB 4t 3 li!*V@ !H 7t!H 5t BqCWDEF7q+W+q+ @`IHBB 6t 3 7!IV@ ] H :t] H 8t BqCWDEF>>7+qq+qW+ @`]  HBB 9t 3 >7 N Ct 04B  J Here competition between synapses appears since strengthening some synapses results in threshold increasing meaning that it is harder for others to be strengthenedD 2` ` z<    z  Dt 00B & R  q depends on postsynaptic activity. For instance, the threshold for LTP decreases when postsynaptic activity is low (y ! q!) F  ` @` `  $`$((x    @      B Ft  fDԔ??0PH t 0޽h ? ̙33___PPT10i.DZpN+D=' B= @B + 0   0j(  0x 0 c $Bp  B  00 TA ? ?   B   0 08F  It is a more direct way of enforcing competition Idea is that postsynaptic neuron can only support a certain amount of total synaptic weight so strengthening one leads to weakening others 2 types: subtractive normalisation and multiplicative normalisationF 2`f ` <  f   H 0 0޽h ? ̙33___PPT10i.м+D=' B= @B + 0  @h(  @x @ c $lF p  F  @0 TA ? ?- "` H $D0 B  @ 0F   * It is easy to prove that the total increase in the weights is 0. FC 2`& ` <  &     @0 TA ? ?` HL   F  @ Z8??C"? @H @ 0޽h ? ̙33___PPT10.0AT@+ !Db' B= @B D' = @BA?%,( < +O%,( < +DT' =%(%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*@%(+ 0 #0(  r  S 4B`  B 4  S A&à400px-BCM_KirkwoodFigure 4: Experimental verification of the BCM rule from Kirkwood et al., 1996.  c0"  Z F ??@ ``  The field potentials evoked in layer III by layer IV stimulation in slices of visual cortex prepared for light-deprived and control rats 4-6 weeks old The effects can be reserved by as little as two days of light exposure before slice preparationFc`_cC<   _   0|3Ffp  ;}Evidence: It is easier to obtain LTP in the cortex of dark-reared animals and it is harder to induced LTD in these cortices j~  c? c 1 0  ? 3H  0޽h ? ̙3380___PPT10.ر: 0 2*@  (   x   c $AF  F    Z??C"?`0P   Z??C"?`P P   ZCF ??P`  , Low- and high-frequency BLA stimuli (LFS, HFS) are known to, respectively, produce homosynaptic NMDA dependent LTD and LTP in ITC cells. Whether LFS and HFS also produce inverse heterosynaptic modifications is unclear. ,U VH   0޽h ? ̙3380___PPT10."08#  0 P#(  v  NA% ??`.   ZTF ??! (Royer and Pare 2003, Nature)gffgf  fgff<   "  Z\1 ?? ~v0___PPT106___PPT9 <" intercalated (ITC) neurons of the amygdala: -N^y~CQ " the basolateral amygdala (BLA): W^YOOgN8h " an array of closely spaced (~150 m) stimulating electrodesc)#cbc` #c" ` #c# # !      "  Zp.\ ??  *n" slices of the amygdala " guinea-pigs (3 5 weeks old)^8``    :  H  0޽h ? ̙3380___PPT10.|Ϊ 0 p(  v  NA) ??& p   ZpQF ??p * p<Plot of EPSP amplitude and rise time versus stimulation site==  ZF ??0  (Royer and Pare 2003, Nature)gffgf  fgff<   J  ZF ??   $" LFS: 0.5 Hz for 10min " HFS consisted of four series of ten trains separated by 0.3 s, each train consisting of ten shocks at 100 Hz " Homosynaptic LTP was induced with HFS paired to postsynaptic depolarization. Postsynaptic depolarization was achieved by applying short (2ms) depolarizing current pulses (0.2 nA) timed so that BLA-evoked EPSPs would occur just before or during current-evoked spikes:cCC> 9H  0޽h ? ̙3380___PPT10.vJ 0 J(  r  S F0  F v  NA( ??`@4   Z F ??  Left: Difference between pre- and post-LFS response profiles (EPSP amplitudes) for one cell (top) and average of all cells Right:Time course of changes in response amplitude{ )  ZF ??!  (Royer and Pare 2003, Nature)gffgf  fgff<   H  0޽h ? ̙3380___PPT10.x0 0 4,(  r  S Fp  F v  NA+ ??@   fF ??` G (Royer and Pare 2003, Nature),gffgf  fgff<   H  0޽h ? ̙3380___PPT10.yP)8 0 @(8(  ( ( 6ta p>6.___PPT9 TIntroduction Synaptic placticity rules " The basic Hebb rule " The covariance rule " BCM Rule " Non-Hebbian rules " Anti-hebbian rules " Timing-based Rules &  Ggc c cccc c $$ $$(g(,G,0g0 4g48g86.___PPT9 (Introduction Synaptic placticity rules " The basic Hebb rule " The covariance rule " BCM Rule " Non-Hebbian rules " Anti-hebbian rules " Timing-based Rules   Ggc c cccc c $$ $$(g(,G,0g0 4g48g8+ !Db' a= @B D' = @BA?%,( < +O%,( < +DT' =%(%(D' =%(D' =4@BBBB%(D' =1:Bvisible*o3>+B#style.visibility<*%(+@0 @(     f\ ?? " About H(): a function like the solid line in previous figure. 6B7> 2   0  `A. ?A ?S"   L  \v  NA ??L@ t H  0޽h ? ̙3380___PPT10.²2  0 6.(  r  S x   v  NA4 ??O/   ZH a ?? `j " The Timing-Based plasticity rule is applied throughout a training period during which the stimulus being presented moves to the right and excites the different neurons in the network sequentially " After the training period, the neuron with sa = 0 receives strengthened input from the sa ="2 neuron and weakened input from the neuron with sa = 2]cC GCC)CCKC3CCKCC>*4H  0޽h ? ̙3380___PPT10.u 0 @8(  v  NA5 ??a   Z85 ?? P. " If the same time-dependent stimulus is presented again after training, the neuron with sa = 0 will respond earlier than it did prior to training " The training experience causes neurons to develop responses that predict the behavior of the stimulus\aWH{%};TYH  0޽h ? ̙3380___PPT10.?[ 0 IA (   r   S ,AP   v   NA7 ?? v   NA8 ??` _%     f8C ??9`  K(Mehta et al. 1997; 2000)B    fG ??} w  h" Place field is negatively skewed after experience(5c2H   0޽h ? ̙3380___PPT10.'+X 0 X(  r  S P    X  0A0?d t X  0A2? D   `t[ ??`  b" Different cortical regions, such as hippocampus and visual cortex have somewhat different forms of synaptic plasticity. " Even in the same cortical region, different types of cells, or even cells within different layers, or even synapses on the same cell that connect to different types of cells, might have varying forms of synaptic plasticity *_c]Cc0&1c06C   f] ??0  V (Abbott and Nelson 2000, Nature)!!fH  0޽h ? ̙3380___PPT10.@Op 0 +#` (  r  S e      Zg ??   kLong-term plastic changes can be induced in about 1 s or less (i.e. within a rather short period, similar to short-term plasticity) The induced change in synaptic weight typically lasts for hours (if no further changes are induced) Changes in synaptic strength are important for formation of memory The longest-lasting forms appear to require protein synthesisH  0޽h ? ̙3380___PPT10.< 0 P  `(  `x ` c $ r    ` ZXs ??@P  >" Unsupervised (or sometimes self-supervised) learning. A network responds to a series of inputs during training solely on the basis of its intrinsic connections and dynamics. The network then self-organizes in a manner that depends on the synaptic plasticity rule being applied and on the nature of inputs presented during training. "a~H ` 0޽h ? ̙33___PPT10i. kg-+D=' = @B + 0 ZR`  d(  d  d Z ??`  (" Supervised learning A desired set of input-output relationships is imposed on the network by a  teacher during training. Networks that perform particular tasks can be constructed in this way Usually, finding a biological plausible teaching mechanism may not be a concern In more biologically plausible examples of supervised learning, one network can act as the teacher for another network"aH d 0޽h ? ̙3380___PPT10. k( 0 ((    Z ??p@  fR" Reinforcement learning It is somewhat intermediate between these cases. The network output is not constrained by a teacher, but evaluative feedback on network performance is provided in the form of reward or punishment H  0޽h ? ̙3380___PPT10.Ȳ 0 p h(  hr h S 8  8 H h 0޽h ? ̙3380___PPT10.k aT 0  l(  lr l S T   H l 0޽h ? ̙3380___PPT10.k0g 0 [S  p(  px p c $    p0 NA ? ?      p0 TA "? ?N0 | "  p Z욚 ?? 0  I5the firing rates of the pre- and postsynaptic neuronsB  p  `D?? B  p  `D??` F p Z8??  `Even with saturation, the basic Hebb rule fails to induce competition between different synapsesa` = p0 TA U? ?@ U H p 0޽h ? ̙3380___PPT10.k0Pa 0  |(  |r | S    H | 0޽h ? ̙3380___PPT10./kzT 0  PT(  Pr P S ح 0    P Z??C"?` H P 0޽h ? ̙3380___PPT10.d0A 0 "(   q B       fд ??0* F(Zhang et al., 1998)H  0޽h ? ̙33y___PPT10Y+D=' = @B + 0  T(  Tr T S ȶ    H T 0޽h ? ̙3380___PPT10.d Y 0 ` Y(    6ǚ  JDo you remember?RB  s *)? @dB  <Ԕ?@  dB  <Ԕ?   dB  <Ԕ?   dB   <Ԕ?  `   <̚r |w12j$      <`Ӛ0pT w Sy"J     <Dؚd7 Eb     BA ?1?oX $0c02   `??02   `??02   `??p0 2   `??0  Z??B  3 rDԔ??PP   fݚ ??P 0p mActivation function"` B   fD>??<  <  |w22j$     <l @G  wn2j(     <TW  Tw3"J     `(xaxa1?0u rx2"    hh88.   `xaxa1?P x16 j $   hh882   `,xaxa1? xn6 j (   hh88B   `D!>??L2  Z??     f ??`   hsimplification"`   BA ?1 ? X $0c0H  0޽h ? ̙3380___PPT10.R 5 0 0 5(  r  S  "0`   v  NA ??PP~v  NA ?? X v  NA ??@ 0    Z$ ??   ?which is positive except in the trivial case when v is constant$@2 H  0޽h ? ̙3380___PPT10.0MX  0 f^P  d(  d d 0/`  ~Note that as keeps changing we need to keep updating the postsynaptic threshold while the presynaptic one is independent of the weightsR 2 `] ` `b   M       f d s *A ? ?  f d s *A ? ? @   d s *A ? ? 8 $D0 d s *A ? ? x8 $D0  d  f= ??  O(homosynaptic depression)    d ZPB ??  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