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How Do Retina Cope With Time Delay?
Chuan Yu Hu
PI: Dr. Chi-Keung Chan
Institute of Physics, Academia Sinica
2020.8.24
1
NPAS Presentation
Introduction
Stimulus and Analysis method
Retina feedback model
Simulation result
Conclusion
Acknowledgement
Reference
Appendix
Outline
2
NPAS Presentation
Introduction | Neural signal has delay effect
Retina inner structure illustration
3
NPAS Presentation
โ€ข There is a time delay in the visual system during the neuron transmission
โ€ข In order to compensate with this delay, a retina need to predict the future
Schnapf, J. L., Kraft, T. W., & Baylor, D. A. (1987).
Inner Retina
Outer Retina
(RGC)
1. Time Delay (30-100ms )
2. Horizontal feedback
3. Parallel processing
NPAS Presentation
4
Stimulus and Analysis method
Time-lag mutual information โ€“ MI(๐œน๐’•)
Response is โ€œpredictiveโ€ if the time lag of
the peak of MI is positive.
๐‘€๐ผ(๐›ฟ๐‘ก๐‘๐‘’๐‘Ž๐‘˜)
r(t)
S(tโ€™)
Future
Past
S(t+ฮ”๐‘ก)
r(t)
Light intensity
๐‘ฅ๐‘ก+ฮ”๐‘ก = (1 โˆ’ ฮ”๐‘ก/๐œ)๐‘ฅ๐‘ก+๐œ‰๐‘ก ๐ทฮ”๐‘ก
Ornstein-Uhlenbeck process (OU)
MI
(
)
NPAS Presentation 5
Retina feedback model
โ€ข Researchers have found different effects by controlling feedback within cone cell โ€“ horizontal cell
โ€ข Derived from experimental result, they have also built models for retina system
Drinnenberg, A. et al. How diverse retinal functions arise from feedback at the first visual synapse. Neuron 99, 117โ€“134.e11 (2018).
Bipolar
Photoreceptor
Horizontal
cell
๐‘ฅ(๐‘ก)
๐‘ฆ(๐‘ก)
๐‘ง(๐‘ก)
Schematic of the model
NPAS Presentation 6
Simulation result of Retina feedback model
Drinnenberg, A. et al. How diverse retinal functions arise from feedback at
the first visual synapse. Neuron 99, 117โ€“134.e11 (2018).
TimeShift (s)
MI
(Arbitrary
Unit)
Input stimulus
MI between cone output and input :
NPAS Presentation 7
Simulation result of Retina feedback model for different ganglion cell type
RGC output
Simulation
Experiment
Predictive cell
Non-Predictive cell
NPAS Presentation 8
Conclusion
1. Cone cell cannot predict without horizontal cellโ€™s feedback
2. Some types of RGC have predictive property
Bipolar
Photoreceptor
Horizontal
cell
๐‘ฅ(๐‘ก)
๐‘ฆ(๐‘ก)
๐‘ง(๐‘ก)
Schematic of the model
NPAS Presentation 9
Reference
[1] Drinnenberg, A. et al. โ€œHow diverse retinal functions arise from feedback at the first visual synapse.โ€ Neuron 99,
117โ€“134.e11 (2018).
[2] Chen, Kevin Sean, Chun-Chung Chen, and C. K. Chan. "Characterization of predictive behavior of a retina by
mutual information." Frontiers in computational neuroscience 11 (2017): 66.
[3] Voss, Henning U. "Signal prediction by anticipatory relaxation dynamics." Physical Review E 93.3 (2016): 030201.
NPAS Presentation 10
End
Questions and advices are welcomed
NPAS Presentation 11
๐‘ฅ๐‘ก+ฮ”๐‘ก = (1 โˆ’ ฮ”๐‘ก/๐œ)๐‘ฅ๐‘ก+๐œ‰๐‘ก ๐ทฮ”๐‘ก
Ornstein-Uhlenbeck process (OU)
This is a Brownian-like walker but this walker cannot move far from the origin. Here ๐œ is
a time constant which governs the correlation time of the walker.
This OU signal is then low-pass filtered with different cut-off frequencies. The filtered
signals become smoother than the original one.
Appendix 1
NPAS Presentation 12
Appendix 2
NPAS Presentation 13
R(t): cone response
h(t): horizontal cell feedback
Appendix 3
๐‘ฆ ๐‘ก = โˆ’๐›ผ๐‘ฆ ๐‘ก + ๐พ ๐‘ฅ ๐‘ก โˆ’ ๐‘ง ๐‘ก
๐‘ง ๐‘ก = โˆ’๐›ฝ๐‘ง ๐‘ก + ๐‘”๐‘ฆ ๐‘ก
๐‘ฆ ๐‘ก = โˆ’๐›ผ๐‘ฆ ๐‘ก + ๐พ ๐‘ฅ ๐‘ก โˆ’ ๐‘ฆ ๐‘ก โˆ’ ๐œ (Voss, Henning U., 2016)
๐‘ฆ ๐‘ก โˆ’ ๐œ is a delayed negative feedback. It is shown that this model gives rise to prediction because
of the negative group delay (NGD) property the system.
In our case, this delay can come from the inhibition feedback of horizontal cell to the cone cell. To
make the model more structural, the horizontal cell feedback can be describe by a new function ๐‘ง(๐‘ก).
The model becomes:
๐›ผ, ๐›ฝ: relaxation time of ๐‘ฆ ๐‘ก , ๐‘ง(๐‘ก)
๐พ: anticipatory coupling constant
๐‘”: input gain of ๐‘ง(๐‘ก)

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How Do Retina Cope With Time Delay?

  • 1. How Do Retina Cope With Time Delay? Chuan Yu Hu PI: Dr. Chi-Keung Chan Institute of Physics, Academia Sinica 2020.8.24 1 NPAS Presentation
  • 2. Introduction Stimulus and Analysis method Retina feedback model Simulation result Conclusion Acknowledgement Reference Appendix Outline 2 NPAS Presentation
  • 3. Introduction | Neural signal has delay effect Retina inner structure illustration 3 NPAS Presentation โ€ข There is a time delay in the visual system during the neuron transmission โ€ข In order to compensate with this delay, a retina need to predict the future Schnapf, J. L., Kraft, T. W., & Baylor, D. A. (1987). Inner Retina Outer Retina (RGC) 1. Time Delay (30-100ms ) 2. Horizontal feedback 3. Parallel processing
  • 4. NPAS Presentation 4 Stimulus and Analysis method Time-lag mutual information โ€“ MI(๐œน๐’•) Response is โ€œpredictiveโ€ if the time lag of the peak of MI is positive. ๐‘€๐ผ(๐›ฟ๐‘ก๐‘๐‘’๐‘Ž๐‘˜) r(t) S(tโ€™) Future Past S(t+ฮ”๐‘ก) r(t) Light intensity ๐‘ฅ๐‘ก+ฮ”๐‘ก = (1 โˆ’ ฮ”๐‘ก/๐œ)๐‘ฅ๐‘ก+๐œ‰๐‘ก ๐ทฮ”๐‘ก Ornstein-Uhlenbeck process (OU) MI ( )
  • 5. NPAS Presentation 5 Retina feedback model โ€ข Researchers have found different effects by controlling feedback within cone cell โ€“ horizontal cell โ€ข Derived from experimental result, they have also built models for retina system Drinnenberg, A. et al. How diverse retinal functions arise from feedback at the first visual synapse. Neuron 99, 117โ€“134.e11 (2018). Bipolar Photoreceptor Horizontal cell ๐‘ฅ(๐‘ก) ๐‘ฆ(๐‘ก) ๐‘ง(๐‘ก) Schematic of the model
  • 6. NPAS Presentation 6 Simulation result of Retina feedback model Drinnenberg, A. et al. How diverse retinal functions arise from feedback at the first visual synapse. Neuron 99, 117โ€“134.e11 (2018). TimeShift (s) MI (Arbitrary Unit) Input stimulus MI between cone output and input :
  • 7. NPAS Presentation 7 Simulation result of Retina feedback model for different ganglion cell type RGC output Simulation Experiment Predictive cell Non-Predictive cell
  • 8. NPAS Presentation 8 Conclusion 1. Cone cell cannot predict without horizontal cellโ€™s feedback 2. Some types of RGC have predictive property Bipolar Photoreceptor Horizontal cell ๐‘ฅ(๐‘ก) ๐‘ฆ(๐‘ก) ๐‘ง(๐‘ก) Schematic of the model
  • 9. NPAS Presentation 9 Reference [1] Drinnenberg, A. et al. โ€œHow diverse retinal functions arise from feedback at the first visual synapse.โ€ Neuron 99, 117โ€“134.e11 (2018). [2] Chen, Kevin Sean, Chun-Chung Chen, and C. K. Chan. "Characterization of predictive behavior of a retina by mutual information." Frontiers in computational neuroscience 11 (2017): 66. [3] Voss, Henning U. "Signal prediction by anticipatory relaxation dynamics." Physical Review E 93.3 (2016): 030201.
  • 10. NPAS Presentation 10 End Questions and advices are welcomed
  • 11. NPAS Presentation 11 ๐‘ฅ๐‘ก+ฮ”๐‘ก = (1 โˆ’ ฮ”๐‘ก/๐œ)๐‘ฅ๐‘ก+๐œ‰๐‘ก ๐ทฮ”๐‘ก Ornstein-Uhlenbeck process (OU) This is a Brownian-like walker but this walker cannot move far from the origin. Here ๐œ is a time constant which governs the correlation time of the walker. This OU signal is then low-pass filtered with different cut-off frequencies. The filtered signals become smoother than the original one. Appendix 1
  • 13. NPAS Presentation 13 R(t): cone response h(t): horizontal cell feedback Appendix 3 ๐‘ฆ ๐‘ก = โˆ’๐›ผ๐‘ฆ ๐‘ก + ๐พ ๐‘ฅ ๐‘ก โˆ’ ๐‘ง ๐‘ก ๐‘ง ๐‘ก = โˆ’๐›ฝ๐‘ง ๐‘ก + ๐‘”๐‘ฆ ๐‘ก ๐‘ฆ ๐‘ก = โˆ’๐›ผ๐‘ฆ ๐‘ก + ๐พ ๐‘ฅ ๐‘ก โˆ’ ๐‘ฆ ๐‘ก โˆ’ ๐œ (Voss, Henning U., 2016) ๐‘ฆ ๐‘ก โˆ’ ๐œ is a delayed negative feedback. It is shown that this model gives rise to prediction because of the negative group delay (NGD) property the system. In our case, this delay can come from the inhibition feedback of horizontal cell to the cone cell. To make the model more structural, the horizontal cell feedback can be describe by a new function ๐‘ง(๐‘ก). The model becomes: ๐›ผ, ๐›ฝ: relaxation time of ๐‘ฆ ๐‘ก , ๐‘ง(๐‘ก) ๐พ: anticipatory coupling constant ๐‘”: input gain of ๐‘ง(๐‘ก)