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Toward a realistic input for visual cortex models
                                                                                                                * Pierre Kornprobst, Serge Picaud*
                                                                                                                      +                                            +                                                                                                                                    +
                                                                   Hassan Nasser, Bruno Cessac, Bogdan Kolomiets,
                                                                                                                                    + NeuroMathComp project team (INRIA, ENS Paris, UNSA, LJAD)                                                                                                                                       Contact: Hassan.Nasser@inria.fr
                                                                                                                                    * Institut de la Vision                                                                                                                                                      http://www-sop.inria.fr/members/Hassan.Nasser
The  modelling  of  the  visual  system  needs  to  provide  realistic  retinal  responses  (spike  trains)  to 
visual  stimuli.  Our  team  [1]  developed  a  retina  simulator  that    allows  a  large  scale  simulation                                       Performing statistics                                                                                                  Simulation of biological experiments
(about 100.000 cells) and provides  spikes train output as a response to visual stimuli. This model 
contains  several  blocks  representing  the  retina  layers.  It  is  able  to  reproduce    realistic  individual 
                                                                                                                                   A  spike  train  is  represented  by  a  set  of  Dirac 
                                                                                                                                
                                                                                                                             functions for  N neurons:
                                                                                                                                                                                                                                                                                      in VirtualRetina
responses  of retinal ganglion cell (RGC). However, as emphasized in the present work, we checked                                                                                                                                                           We simulated the biological experiment (described before) with VirtualRetina. A 93 sec of image sequence. The spontaneous 




                                                                                                                                                                                              Neuron Index
that the collective responses of RGC don't match to real data. Comparing our simulator outputs to                                                                                                                                                         activity is simulated with a white noise (dark image), the light flash stimulus with a white noise (lighted image). We respected 
real acquisitions data made by  B. Kolomiets  and S. Picaud (Institut de la Vision)  we have shown,                                                                                                                                                       the exact time of stimulus (1 sec of light flash followed by 5 sec of rest repeated 10 times) in order to reproduce the experiment 
                                                                                                                                 We call observables, the events we observe in this                                                                       conditions. Fig. 11 and 12 show C(T) respectively in spontaneous and evoked potential activity.
using  statistical  methods  developed  by  our  team  (EnaS)  [3],    that  real  data  show  a  significant                spike  train  (Ex:  Individual  spike  related  with  firing 
synchronization and correlations between RGC outputs, that is absent from our simulator outputs.                             rate  and  doublets  related  with  correlations  ...).  The                                                                                                                       Simulation  with  the  VirtualRetina 
                                                                                                                                                                                                                                                                                                                software  shows  a  decreasing  of  the 
It  is  commonly  believed  that  those  correlations  come  from  the  overlap  of  RGC  receptive  fields.                 correlation is given by the following equation:
                                                                                                                                                                                                                                                                                                                correlation  with  the  raster  length  (Both 
However, since, our simulator carefully reproduces this overlap this suggests that synchronization                                                                                                                                            Time (s)
                                                                                                                                                                                                                                                                                                                in  spontaneous  and  evoked  potential 
                                                                                                                                                                                                                           Fig. 3
must be explained by other mechanisms such as long range connections from Amacrine to RGC or                                                                                                                                                                                                                    activities).  The  synthetic  data  (two 
electrical connections (gap junctions) between RGC which have not been implemented yet in  our                               Where the first term represents how many times the neurons i fires after a time delay t from the                                                                                   neurons  firing  independently)  show  also 
simulator.                                                                                                                   neuron j.     is the empirical average of an observable.                                                                                                                           the  same  behavior.  This  is  the  typical 
                                                                                                                                                                                                                                                                                                                behavior  of  two  non-correlated  firing 
                                                                                                                                  We consider the evolving of the correlation value in                                                                                                                          neurons.  Despite  the  fact  that  the 
                Problematic & Background                                                                                     term  of  the  raster  length  (T)  in  order  to  evaluate  the 
                                                                                                                             correlation  between  two  neurons.  Theoretically,  this 
                                                                                                                                                                                                                                                                                                                receptive  fields  are  implemented  in  the 
                                                                                                                                                                                                                                                                                                                software, the correlation outlook for two 
                                                                                                                             correlation tends to zero when T tends to infinity (Plot                                                                                                                           neurons  in  the  software  reflects  the 
                                                      Spike Train                                                            in the log scale) as    /       , Where K is a constant.                                                                                                                           behavior  of  non-correlated  neurons.  Our 
        Stimulus                   Retina                                       Visual                     Cortex                                                                                                                                                                                               first  conclusion  is  that  the    overlap  of 
                                                                                Cortex                     Response                                                                                                                                                                                             RGC  receptive  fields  in  Virtual  Retina   
                                                                                                                             Fig.  4  shows  C(T)  for  two  neurons  that  fire 
                                                  Retinal        Cortex                                                                                                                                                                                                                                         doesn't imply correlation between RGC.                               Fig. 12
                                                                                                                             independently.  Despite  the  fluctuations  on  small  time                                                                                       Fig. 11
                                                  output         Input
                                                                                                                             windows, we see the evolution of C(T):
      The modeling of the visual system needs to provide realistic retinal responses (spike trains) to 
  visual stimuli. Our team (Wohrer et al. 2008) developed a retina simulator that  allows a large scale 
  simulation (about 100.000 cells) and provides spikes train outputs as a response to visual stimuli. 
                                                                                                                                                                                                                           Fig. 4    Raster length (s)
                                                                                                                                                                                                                                                                                       Interaction between RGC cells
  This model contains several blocks representing the retina layers.                                                                                                                                                                                     In order to have hints on the RGC circuitry, we                                                                       Fig.  14  shows  the      for 
          Biological  experiments  are  time  consuming  and  expensive.  An  alternative  could  be  a  retina 
  simulator: The VirtualRetina.
                                                                                                                                                   Statistics on real data                                                                               applied  Ising  model  to  analyze  statistics  of 
                                                                                                                                                                                                                                                         these  data.  Ising  model  is  characterized  by  a 
                                                                                                                                                                                                                                                                                                                                                                               neurons  at  several  distance. 
                                                                                                                                                                                                                                                                                                                                                                               The  units  are  the  indexes  of 
                                                     Retinal connection [4]     VirtualRetina model [1]                                                                                                                               Fig. 5
  The vertebrate retina                                                                                                      Real  data  acquisition  were  provided  by  Bogdan                                                                         Gibbs distribution whose potential is:                                                                                neurons  in  the  MEA  chip. 
                                                                                                                             Kolmiets and Serge Picaud (Institut de la vision de                                                                                                                                                                                               L0LX  coefficients  are  for        . 
                                                                                                                             Paris).  Acquisition  details:  30  sec  of  spontaneous                                                                                                                                                                                          The  index  0  replaces  the 
                                                                                                                             activity  followed  by  63  sec  of  evoked  potential  (1                                                                                                                                                                                        unit  8,  the  index  X  replaces 
                                                                                                                                                                                                                                                         Ising  model  is  one  of  the  most  popular  in 
                                                                                                                             sec of light flash followed by 5 sec of rest, repeated                                                                      neural  computation.  It  is  believed  that  the                                                                     the  other  units:  16,  24,  32, 
                                                                                                                             several  10  times)  -  Acquisition  on  MEA  chip  of  54                                                                  coefficients    are  related  to  connections                                                                         39,  47.  These  units  are 
                                                                                                                             electrodes.  The  activity  of  these  neurons  is  shown                                                                   between  gap  junctions  although  other                                                                              located  at  various  distances 
                                                                                                                                                                                                                                                         interpretations are possible.                                                                              Fig. 13
                                                                                                                             in the Fig. 5.                                                                                                                                                                                                                                    from  the  unit  0.    the 
                                                                                                                                                                                                                                                         Fig. 13 shows the     and      for a two different                                                                    coefficient      increases  and 


                                                                 Fig. 2                           Fig. 3
                                                                                                                             Experiments:                                                                                                                sets of 7 neurons (N0-N6). The    are the firing 
                                                                                                                                                                                                                                                         rate  (in  positive).  The          are  gives  an  idea 
                                                                                                                                                                                                                                                                                                                                                                               then  decrease  after  a 
                                                                                                                                                                                                                                                                                                                                                                               distance  of  about  300  um. 
                          Fig. 1                                                                                                                                                                                                                         about  the  RGC  interaction.  Interaction  values                                                                    How  can  we  interpret  this 
                                                                                The  diagram  of  the  VirtualRetina.        Experiments were done for rats' retina, aged between 13 and 17 months. The electrode (Fig.                                  appear almost in negative. Knowing that RGC                                                                           bahavior?
   The  retina  contains  several  layer  of         Connectivity  map                                                       6) size is: 40x40 um. The inter-electrode distance is 200 um. the retina were placed in a
                                                                                The  receptors  and  horizontal  layer                                                                                                                                   are connected with Gap Juction which implies 
   cells  connected  through  chemical               in  the  retina.  full 
                                                                                are  implemented  as  image  filters.        recording chamber where the MEA and stimulation tools are (Fig. 7).                                                         a positive connectivity. Why do these negative 
   and electrical synapse. More than 50              and  empty  disks          However,  bipolar  and  ganglion                                                                                                                                                                                                                                                              Fig. 14
                                                     represent                                                                                                                                                                                           values appear?
   cell  subtypes  exists.  However,                                            cells     are    implemented        as                                        Fig. 6
   scientist believe that less than 50 %             respectively               conductance  based  and  I&F 
   of  cell  functions  are  known  until            excitatory       and       neurons  respectively.  Amacrine 
   now.                                              inhibitory.                cells are not yet implemented.                                                                                               Fig. 7
                                                                                                                                                                                                                                                                                                                     Discussion
                                                                                                                                                                                                                                                                            Fig. 15 [5]
         Performance of VirtualRetina                                                                                                                                                                                                                                                                                                                      Fig.  15  shows  cros  correlograms  for  directely 
                                                                                                                                                                                                                                                                                                                                                           connected  RGC  and  Amacrine  connected  RGC. 
          Allows large scale simulation (More than 100,000 cells).                                                                                                                                                                                                                                                                                         The  direct  connection  implies  a  narrow 
          Possibility of customizing retina and retina parameters.                                                                                                                                            Fig. 8 [2]                                                                                                                                   correlation.  In  contrary,  broad  correlation 

  +       High biological plausibility at the level of single cell.
          Reasonable computational cost.
                                                                                                                                                                                                                                                                                                                                                           appears  for  RGC  connected  through  Amacrine 
                                                                                                                                                                                                                                                                                                                                                           cells.
          Implements the underlying of receptive fields of retinal ganglion cells (RGC).

          Amacrine cells are not implemented.                                                                                                                                                                                                                     The  VirtualRetina  needs  to  be  improved  at  the  level  of  RGC  cells  circuitry.  One  of  our  perspectives  is  to  implement  gap 
    -     Connections between RGC and Amacrine-RGC are not implemented.
          Statistically, the response of a set of RGC doesn't fit to real RGC.
                                                                                                                                                                                                                                                          junction connections between RGC in order to approach statistics on real data acquisition.
                                                                                                                                                                                                                                                               Amacrine cells provide a negative feedback to RGC cells. The negative connectivity coefficients could be interpreted by these 
                                                                                                                                                                                                                                                          feedback.
                                                                                                                                                                                                                                                               A statistically plausible retinal model is required to produce a realistic retinal input for the visual cortex model.
                                                                 [3] J.C. Vasquez, T. Vieville, B. Cessac, Entropy-based                                                  Fig. 9                                                    Fig. 10                   We would like to create image-statistics dictionary. In fact, we believe that the response of the RGC depends statistically on 
  [1] Adrien Wohrer, Model and large-scale simulator of a                                                                                                                                                                                                 the image events (motion, color detection, texture detection., ....). The implementation of retinal circuitry provided in (Gollish et 
                                                                 parametric estimation of spike train statistics. INRIA,
  biological retina, with contrast gain control. University of
  Nice Sophia-Antipolis, INRIA, 2009.
                                                                 2009.                                                                                                                                                                                    al. 2009) will help to create this dictionary.
                                                                 [4] T. Gollisch, M. Meister. Eye Smarter than Scientists
  [2] B. Kolomiets, E. Dubus, M Simonutti, S Rosolen, J.A        Believed: Neural Computations in Circuits of the Retina.
                                                                                                                             Fig.  9  and  10  show  C(T)  in  spontaneous  activity  and  evoked  potential  respectively,  for  three                     
  Sahel, S.Picaud, Late his-tological and functional changes
  in the P23H rat retina afterphotoreceptor loss.
                                                                 j.neuron. 2009 (volume 65 issue 2 pp.150 - 164) .           different pairs of neurons mounted at several distances. The correlation outlook is different than                                                                                            Softwares
  Neurobiology of Disease 38 (2010) 47-58.
                                                                 [5] S. Bloomfield, B. Völgyi. The diverse functional roles   for independently firing neurons. This assumption implies that there exist connections within the                           VirtualRetina: http://www-sop.inria.fr/neuromathcomp/software/virtualretina/index.shtml
                                                                 and regulation of neuronal gap junctions in the retina.
                                                                                                                             network of RGC which induces synchronization in firing.
                                                                 Nature Reviews Neuroscience 10, 495-506 (July 2009).                                                                                                                                    EnaS (Event neural assembly simulator): http://enas.gforge.inria.fr/
                                                                                                                                                                                                                                                         .


                                                                                                                  http://www-sop.inria.fr/neuromathcomp
Toward a realistic input for visual cortex models: Simulating retinal correlations

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Toward a realistic input for visual cortex models: Simulating retinal correlations

  • 1. Toward a realistic input for visual cortex models * Pierre Kornprobst, Serge Picaud* + + + Hassan Nasser, Bruno Cessac, Bogdan Kolomiets, + NeuroMathComp project team (INRIA, ENS Paris, UNSA, LJAD) Contact: Hassan.Nasser@inria.fr * Institut de la Vision http://www-sop.inria.fr/members/Hassan.Nasser The  modelling  of  the  visual  system  needs  to  provide  realistic  retinal  responses  (spike  trains)  to  visual  stimuli.  Our  team  [1]  developed  a  retina  simulator  that    allows  a  large  scale  simulation  Performing statistics Simulation of biological experiments (about 100.000 cells) and provides  spikes train output as a response to visual stimuli. This model  contains  several  blocks  representing  the  retina  layers.  It  is  able  to  reproduce    realistic  individual        A  spike  train  is  represented  by  a  set  of  Dirac    functions for  N neurons: in VirtualRetina responses  of retinal ganglion cell (RGC). However, as emphasized in the present work, we checked    We simulated the biological experiment (described before) with VirtualRetina. A 93 sec of image sequence. The spontaneous  Neuron Index that the collective responses of RGC don't match to real data. Comparing our simulator outputs to  activity is simulated with a white noise (dark image), the light flash stimulus with a white noise (lighted image). We respected  real acquisitions data made by  B. Kolomiets  and S. Picaud (Institut de la Vision)  we have shown,  the exact time of stimulus (1 sec of light flash followed by 5 sec of rest repeated 10 times) in order to reproduce the experiment      We call observables, the events we observe in this  conditions. Fig. 11 and 12 show C(T) respectively in spontaneous and evoked potential activity. using  statistical  methods  developed  by  our  team  (EnaS)  [3],    that  real  data  show  a  significant  spike  train  (Ex:  Individual  spike  related  with  firing  synchronization and correlations between RGC outputs, that is absent from our simulator outputs.  rate  and  doublets  related  with  correlations  ...).  The  Simulation  with  the  VirtualRetina  software  shows  a  decreasing  of  the  It  is  commonly  believed  that  those  correlations  come  from  the  overlap  of  RGC  receptive  fields.  correlation is given by the following equation: correlation  with  the  raster  length  (Both  However, since, our simulator carefully reproduces this overlap this suggests that synchronization  Time (s) in  spontaneous  and  evoked  potential  Fig. 3 must be explained by other mechanisms such as long range connections from Amacrine to RGC or  activities).  The  synthetic  data  (two  electrical connections (gap junctions) between RGC which have not been implemented yet in  our  Where the first term represents how many times the neurons i fires after a time delay t from the  neurons  firing  independently)  show  also  simulator. neuron j.     is the empirical average of an observable. the  same  behavior.  This  is  the  typical  behavior  of  two  non-correlated  firing       We consider the evolving of the correlation value in  neurons.  Despite  the  fact  that  the  Problematic & Background term  of  the  raster  length  (T)  in  order  to  evaluate  the  correlation  between  two  neurons.  Theoretically,  this  receptive  fields  are  implemented  in  the  software, the correlation outlook for two  correlation tends to zero when T tends to infinity (Plot  neurons  in  the  software  reflects  the  Spike Train in the log scale) as    /       , Where K is a constant. behavior  of  non-correlated  neurons.  Our  Stimulus Retina Visual Cortex  first  conclusion  is  that  the    overlap  of  Cortex  Response RGC  receptive  fields  in  Virtual  Retina    Fig.  4  shows  C(T)  for  two  neurons  that  fire  Retinal  Cortex doesn't imply correlation between RGC. Fig. 12 independently.  Despite  the  fluctuations  on  small  time  Fig. 11 output Input windows, we see the evolution of C(T):     The modeling of the visual system needs to provide realistic retinal responses (spike trains) to  visual stimuli. Our team (Wohrer et al. 2008) developed a retina simulator that  allows a large scale  simulation (about 100.000 cells) and provides spikes train outputs as a response to visual stimuli.  Fig. 4 Raster length (s) Interaction between RGC cells This model contains several blocks representing the retina layers. In order to have hints on the RGC circuitry, we  Fig.  14  shows  the      for          Biological  experiments  are  time  consuming  and  expensive.  An  alternative  could  be  a  retina  simulator: The VirtualRetina. Statistics on real data applied  Ising  model  to  analyze  statistics  of  these  data.  Ising  model  is  characterized  by  a  neurons  at  several  distance.  The  units  are  the  indexes  of  Retinal connection [4] VirtualRetina model [1] Fig. 5 The vertebrate retina Real  data  acquisition  were  provided  by  Bogdan  Gibbs distribution whose potential is: neurons  in  the  MEA  chip.  Kolmiets and Serge Picaud (Institut de la vision de  L0LX  coefficients  are  for        .  Paris).  Acquisition  details:  30  sec  of  spontaneous  The  index  0  replaces  the  activity  followed  by  63  sec  of  evoked  potential  (1  unit  8,  the  index  X  replaces  Ising  model  is  one  of  the  most  popular  in  sec of light flash followed by 5 sec of rest, repeated  neural  computation.  It  is  believed  that  the  the  other  units:  16,  24,  32,  several  10  times)  -  Acquisition  on  MEA  chip  of  54  coefficients    are  related  to  connections  39,  47.  These  units  are  electrodes.  The  activity  of  these  neurons  is  shown  between  gap  junctions  although  other  located  at  various  distances  interpretations are possible. Fig. 13 in the Fig. 5. from  the  unit  0.    the  Fig. 13 shows the     and      for a two different  coefficient      increases  and  Fig. 2 Fig. 3 Experiments: sets of 7 neurons (N0-N6). The    are the firing  rate  (in  positive).  The          are  gives  an  idea  then  decrease  after  a  distance  of  about  300  um.  Fig. 1 about  the  RGC  interaction.  Interaction  values  How  can  we  interpret  this  The  diagram  of  the  VirtualRetina.  Experiments were done for rats' retina, aged between 13 and 17 months. The electrode (Fig. appear almost in negative. Knowing that RGC  bahavior? The  retina  contains  several  layer  of  Connectivity  map  6) size is: 40x40 um. The inter-electrode distance is 200 um. the retina were placed in a The  receptors  and  horizontal  layer  are connected with Gap Juction which implies  cells  connected  through  chemical  in  the  retina.  full  are  implemented  as  image  filters.  recording chamber where the MEA and stimulation tools are (Fig. 7). a positive connectivity. Why do these negative  and electrical synapse. More than 50  and  empty  disks  However,  bipolar  and  ganglion  Fig. 14 represent  values appear? cell  subtypes  exists.  However,  cells  are  implemented  as  Fig. 6 scientist believe that less than 50 %  respectively  conductance  based  and  I&F  of  cell  functions  are  known  until  excitatory  and  neurons  respectively.  Amacrine  now. inhibitory. cells are not yet implemented. Fig. 7 Discussion Fig. 15 [5] Performance of VirtualRetina Fig.  15  shows  cros  correlograms  for  directely  connected  RGC  and  Amacrine  connected  RGC.    Allows large scale simulation (More than 100,000 cells). The  direct  connection  implies  a  narrow    Possibility of customizing retina and retina parameters. Fig. 8 [2] correlation.  In  contrary,  broad  correlation  +   High biological plausibility at the level of single cell.   Reasonable computational cost. appears  for  RGC  connected  through  Amacrine  cells.   Implements the underlying of receptive fields of retinal ganglion cells (RGC).   Amacrine cells are not implemented.           The  VirtualRetina  needs  to  be  improved  at  the  level  of  RGC  cells  circuitry.  One  of  our  perspectives  is  to  implement  gap  -   Connections between RGC and Amacrine-RGC are not implemented.   Statistically, the response of a set of RGC doesn't fit to real RGC. junction connections between RGC in order to approach statistics on real data acquisition.      Amacrine cells provide a negative feedback to RGC cells. The negative connectivity coefficients could be interpreted by these  feedback.      A statistically plausible retinal model is required to produce a realistic retinal input for the visual cortex model. [3] J.C. Vasquez, T. Vieville, B. Cessac, Entropy-based Fig. 9 Fig. 10     We would like to create image-statistics dictionary. In fact, we believe that the response of the RGC depends statistically on  [1] Adrien Wohrer, Model and large-scale simulator of a the image events (motion, color detection, texture detection., ....). The implementation of retinal circuitry provided in (Gollish et  parametric estimation of spike train statistics. INRIA, biological retina, with contrast gain control. University of Nice Sophia-Antipolis, INRIA, 2009. 2009. al. 2009) will help to create this dictionary. [4] T. Gollisch, M. Meister. Eye Smarter than Scientists [2] B. Kolomiets, E. Dubus, M Simonutti, S Rosolen, J.A Believed: Neural Computations in Circuits of the Retina. Fig.  9  and  10  show  C(T)  in  spontaneous  activity  and  evoked  potential  respectively,  for  three     Sahel, S.Picaud, Late his-tological and functional changes in the P23H rat retina afterphotoreceptor loss. j.neuron. 2009 (volume 65 issue 2 pp.150 - 164) . different pairs of neurons mounted at several distances. The correlation outlook is different than  Softwares Neurobiology of Disease 38 (2010) 47-58. [5] S. Bloomfield, B. Völgyi. The diverse functional roles for independently firing neurons. This assumption implies that there exist connections within the  VirtualRetina: http://www-sop.inria.fr/neuromathcomp/software/virtualretina/index.shtml and regulation of neuronal gap junctions in the retina. network of RGC which induces synchronization in firing. Nature Reviews Neuroscience 10, 495-506 (July 2009). EnaS (Event neural assembly simulator): http://enas.gforge.inria.fr/ . http://www-sop.inria.fr/neuromathcomp