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Georgopoulos s group carried out a series elegant experiments to explore the neural representation of moving direction. :5I|`&l 9 P <( Vt [It is one of the few examples in which the coding scheme is relatively well understood. :3)Z`[ H P 0޽h ? ̙33` c 0L0 og.T (  T ~ T s *'( q  ' P T <P ~In the experiment, the monkey is guided to move the lever in the center of apparatus to one of eight peripheral s. .B=~`~  T <s2 1Neural activities in the motor area are recorded.$21`1  T ZA ( 10_18g 'F T <fM VAll pictures from M. Gazzaniga et al. Cognitive Neuroscience, unless otherwise stated.:W&bbb&   9 H T 0޽h ? ̙33y___PPT10Y+D=' {"= @B + d 0L0  .\ j(  \ ~ \ s *'( w   % \ <x?<q$ Preferred stimulus: for each neuron, there is a stimulus value by which the neuron has the maximum response. Tuning curve: the function mapping between the neural activity (measured by mean firing rate) and the stimulus value. bCL@Q} `  \ <O"'B oNoises always exist$`  \ \A( 10_19u' H \ 0޽h ? ̙33y___PPT10Y+D=' {"= @B + e 0L0 H@@.d (  d x d c $p( Pp     d <v  How is the moving direction encoded by neural activities? By the most active neuron? This sounds reasonably if there is no noise, but it does not work in practice because of large fluctuations in neural activities. By a population of neurons? All active neurons contain a piece of information about the stimulus, why don t we consider all of them jointly encode the moving direction. It has at least one apparent advantage of averaging out noises in individual neurons, since they are (partially) independent. Georgopoulos et al. proposed an idea to reconstruct the moving direction from the observed neural activities.nxc?1 ~`& c H d 0޽h ? ̙33 f 0L0 &`.l (  l x l c $ٸ( @   > l <0P6 ZThe smooth bell-shape tuning curve is often modeled by the Gaussian (or cosin) function: .; Z`&I  ` l  c $A (??@ @  ( l <F . f  | For neural activity in one trial$! `  ` l  c $A )?? ( )H l 0޽h ? ̙33|  0L0  .t (  t x t c $(     t <$p Georgopoulos s idea: the neural system reads out the moving direction by the average of preferred stimuli of all active neurons weighted by their activities. This sounds reasonable since more active neurons, whose preferred stimuli are more likely close to the true stimulus, and hence should contribute more on the final vote..<T`F ` t  c $A *??p *H t 0޽h ? ̙332 g 0L0 .x Z(  x ~ x s *(     x \A ( 10_19lO H x 0޽h ? ̙33  0L0 <4. (   x  c $( `@     <Lr Population vector demonstrated that information can be accurately represented by the joint activities of a population of neurons in a noise environment. This coding strategy of using a population of neurons to represent stimulus is called population coding. BB`   <@m  ?The idea of population coding is also found in the representation of moving direction in other parts of cortex, and the representation of other stimuli, such as the orientation of object and the spatial ..E` k  <pj V  Population coding seems to be a general framework for information processing in neural systems, and worth to be analyzed in more detail theoretically. .BX` H  0޽h ? ̙33 h 0L0 $. (   x  c $3( w     <9z' nThe encoding phase$` `   c $A +??! +`   c $A ,??H  ,  <F P  nThe decoding phase$` `   c $A -??0 k  -  <Y  6Population vector is one of many inference strategies.&76b6 H  0޽h ? ̙33  0L0 JB. 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H @ 0޽h ? ̙3380___PPT10.x`>j 0 .X (  X d X c $    X s * @   xA light comes on at a target position, signalling the moving direction. The monkey is trained to following the guidance in order to get food reward. Since the arm movement of monkey involves the function of motor cortex, Georgopoulos then recorded the neural activities in the motor cortex, and their observation of the activities of a particular neuron is shown in this figure. The neural activity of the same neuron for each possible moving direction is drawn at the corresponding target position. For example. Many trials results are illustrated. A dot represents one spike. Each row represents one Trial. There are two observations are notable. One is that the neuron shows different activity levels, i.e., different Firing rates, when different moving directions are presented. This implies the information of moving direction is In the activities of neurons in the motor area. Secondly, this neuron is not only to be activated by a single moving direction, but by many of them, though the strengths are different. `:'   H X 0޽h ? ̙33Ak 0 0.` (  ` d ` c $    ` s *Y @   {#These two observations are also true for other stimuli instead of moving direction, and motivated neuroscientists to propose two important concepts to describe neural coding. The following figure shows a typical example for the tuning curve of a neuron encoding the moving direction. The left hand figure shows that neural responses when different values of moving direction are presented. This neuron has the maximum response when the stimulus is 180 degree. This means 180 degree is the preferred Stimulus of this neuron. Here we consider the neural activity is measured the firing rate. This firing rate is of cause, fluctuated trial by Trial. Suppose we take its mean value over many trails. The right hand side figure draws the relationship between the mean firing of this neuron and the stimulus. Interestingly, we see that the tuning curve has smooth, symmetric, bell shape. This shape tells us at least two things. Symmetry implies the tuning curve must be a function of the difference between the preferred stimulus and The actual stimulus, instead of a direct function of the actual stimulus. Secondly, the bell shape, means that the Tuning curve is decay function of this difference, I.e., the neural activity decreases with the difference between the stimulus and the neuronal preferred stimulus. $#`# H ` 0޽h ? ̙33l 0 h`P.h (  h d h c $   T h s *x˸ @   In the above, we show the experimental observation. Though we note the correlation between moving directions and the neuronal firing rates, we have not figure out the coding scheme exactly. One simple suggestion, the most active neuron reports the stimulus value, since we know when The Stimulus equals to a neuron s preferred stimulus, this neuron s tuning function has the largest value. However, this does not work well in reality due to the noise. ` H h 0޽h ? ̙33 m 0 p.p Z(  p d p c $    p s * @   DThis Gaussian function captures the main features of neural responses. It has the bell-shape. The neuron I responses most when stimulus x is equal to its preferred stimulus c_I, and decreases with the their difference.`(  H p 0޽h ? ̙33 o 0 P H .|  (  | d | c $   <  | s * @   r Let us first look at the left hand side picture. Consider two neurons with their tuning functions as shown in this figure. They have different preferred-stimuli. Cell 1 has the preferred stimulus 180 degree, cell2 has 90 degree. Suppose the real stimulus is 180 degree. So in most case, celle1 is more active than cell2. The weight summation in population vector will be looks like this. The red vector represents the weighted contribution. Its direction is the cell1 s preferred one, and the length represent the neural activity. Similarly, for the green vector which represents The contribution of cell2. Since the real stimulus is close to the cell1 s preferred stimulus, cell1 tends to be more active, and hence the red vector is longer. As a result, the summed result, will be more towards the direction of Red vector. This is we expected. Similarly for the case when the real stimulus is 90 degree is shown here. In this example, since there is only two neurons the decoding result is not good. If there are many neurons, as is the case in reality, the result will be much better. This picture shows the result of using population vector when working with experimental data. Each brown vector represent the weight contribution of each neuron. You can Individually they are very noise, but the population vector works very well. :9`9 H | 0޽h ? ̙33q 0 vn. 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Nq+The mathematical model of population coding(,+F33+ Or Rg&&0 PsRg&&0 QtRg&&0 RuRg&&0 SvRg&&0 TwRg&&0 Ux TnRg0g0&0 Vy _nRg0g0&0 WzRg&0 X{Rg&0 /<bYcZd[e\f]g^h  0N0 sk-P (  P x P c $I ( P `    j P 0T   s Two pieces of information are important for representing the dynamical aspects of external objects, namely, the moving speed and the moving direction of objects. .@  P <`   >@   P <(d 0 V  |A. Georgopoulos s group carried out a series elegant experiments to explore the neural representation of moving direction. 85I|@&l 7 P <l"  Vt [It is one of the few examples in which the coding scheme is relatively well understood. 83)Z@[ H P 0޽h ? ̙33V c 0N0 e].T (  T ~ T s *y ( q    N T <[Y MS ~In the experiment, the monkey is guided to move the lever in the center of apparatus to one of eight peripheral s. ,B=~@~  T <0 2 1Neural activities in the motor area are recorded."21@1  T ZA ( 10_18g  @ T <  fM VAll pictures from M. Gazzaniga et al. Cognitive Neuroscience, unless otherwise stated.4W&BBB&   9 H T 0޽h ? ̙33y___PPT10Y+D='  = @B + d 0N0  .\ f(  \ ~ \ s *| ( w    # \ <x q$ Preferred stimulus: for each neuron, there is a stimulus value by which the neuron has the maximum response. Tuning curve: the function mapping between the neural activity (measured by mean firing rate) and the stimulus value. `CL@Q} @  \ <tz }' mNoises always exist"@  \ \A( 10_19u'  H \ 0޽h ? ̙33y___PPT10Y+D='  = @B + e 0N0 F>@.d (  d x d c $ ( Pp     d <   How is the moving direction encoded by neural activities? By the most active neuron? This sounds reasonably if there is no noise, but it does not work in practice because of large fluctuations in neural activities. By a population of neurons? All active neurons contain a piece of information about the stimulus, why don t we consider all of them jointly encode the moving direction. It has at least one apparent advantage of averaging out noises in individual neurons, since they are (partially) independent. Georgopoulos et al. proposed an idea to reconstruct the moving direction from the observed neural activities.lxc?1 ~@& c H d 0޽h ? ̙33 f 0N0 "`.l (  l x l c $ĺ ( @    < l <H@ 0P6 ZThe smooth bell-shape tuning curve is often modeled by the Gaussian (or cosin) function: ,; Z@&I  ` l  c $A (??@ @  ( l <E F . f  z For neural activity in one trial"! @  ` l  c $A )?? ( )H l 0޽h ? ̙33z  0N0  .t (  t x t c $ (      t <O M Georgopoulos s idea: the neural system reads out the moving direction by the average of preferred stimuli of all active neurons weighted by their activities. This sounds reasonable since more active neurons, whose preferred stimuli are more likely close to the true stimulus, and hence should contribute more on the final vote.,<T@F ` t  c $A *??p *H t 0޽h ? ̙332 g 0N0 .x Z(  x ~ x s *n ( w    x \A ( 10_19lO  H x 0޽h ? ̙33  0N0 6.. (   x  c $\ ( `@     <\b Mq- Population vector demonstrated that information can be accurately represented by the joint activities of a population of neurons in a noise environment. This coding strategy of using a population of neurons to represent stimulus is called population coding. @B@   <u   =The idea of population coding is also found in the representation of moving direction in other parts of cortex, and the representation of other stimuli, such as the orientation of object and the spatial .,E@ i  <s j V  Population coding seems to be a general framework for information processing in neural systems, and worth to be analyzed in more detail theoretically. ,BX@ H  0޽h ? ̙33 h 0N0 . (   x  c $ ( w     < z' lThe encoding phase"@ `   c $A +??! +`   c $A ,??H  ,  <  P  lThe decoding phase"@ `   c $A -??0 k  -  <Ȏ   6Population vector is one of many inference strategies.$76B6 H  0޽h ? ̙33  0N0 . 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