amol gharat and curtis bakeramolgharat.com/model nonlinear y cell receptive field estimation.pdf ·...

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INTRODUCTION 0.1 1 0.5 2 4 0 2 4 6 8 10 Linear (F1) NonLinear (F2) Spatial Frequency (cpd) Firing Rate (Hz) Y-cell responses to texture stimuli predicted by a nonlinear receptive field model based on bipolar cell subunits 1 Integrated Program in Neuroscience, McGill Univ.; 2 Dept of Ophthalmology, McGill Vision Res. Unit, McGill University, Montreal, Canada Amol Gharat 1 and Curtis Baker 2 METHODS RESULTS CONCLUSIONS 1. Rosenberg et al (2010). Subcortical representation of non-fourier image features. J Neuroscience. 30(6):1985-1993. 2. Demb et al (2001a). Cellular basis for the response to second-order motion cues in Y retinal ganglion cells. Neuron. 32:711-721. 3. Demb et al (2001b). Bipolar cells contribute to nonlinear spatial summation in the brisk-transient (Y) ganglion cell in mammalian retina. J Neuroscience. 21(19):7447-7454. 4. Borghuis et al (2013) Two-photon imaging of nonlinear glutamate release dynamics at bipolar cell synapses in the mouse retina. J Neuroscience. 33(27):10972-10985. 5. Hochstein & Shapley (1976). Linear and nonlinear spatial subunits in Y cat retinal ganglion cells. 262:265-284. 6. http://strflab.berkeley.edu/ Funded by CIHR grant MOP-119498 to CB. Vanier Canada Graduate Scholarship to AG. Many thanks to Guangxing Li and Vargha Talebi. Y cell “subunit” receptive field X cell receptive field Variety of Y cell receptive fields time Y-cell model Model Spatial Tuning 1 2 4 8 0.5 0.25 35 40 45 50 55 Subunit Size VAF % Linear HWR FWR 0 20 40 60 Subunit Rectification Type VAF % - A prime aim of visual neuroscience is to estimate neuronal receptive field models that can predict responses to arbitrary stimuli. - X (linear) and Y (non-linear) type cells form two major categories in mammalian retina and LGN. - Y type cells may be an important early stage for processing second-order information in visual scenes (Demb et al., 2001a; Rosenberg et al., 2010). - A linear receptive field model with static output nonlinearity (LN model) cannot account for nonlinear responses of Y-cells to high resolution texture stimuli. - A model with spatial pooling of rectified inputs from several small subunit receptive fields (Demb et al., 2001b) might better predict responses of Y-cells to arbitrary texture stimuli. Stimuli - Synthetic Naturalistic Texture images at 75Hz - 3 datasets training (7500 images) regularization (1875 images) validation (1875 images) - Sinusoidal gratings (Drifting or Contrast reversing) Animal Preparation - Anesthetized and paralyzed cats - Extracellular, single-unit recordings in LGN using single & multi-channel electrodes - Manual spike sorting using Plexon Offline Sorter & SpikeSorter (UBC) Classification of Cells (X or Y) - Measured responses of neurons to contrast reversing gratings at series of spatial frequencies - If response at second harmonic (F2) was greater than first harmonic (F1), neuron was classified as Y-type Receptive Field Estimation - Preprocessing with Gaussian subunits - Estimation of subunit weights with gradient descent optimization.(STRFlab Software) - Regularization (early stopping) - Measure model performance for series of subunit sizes and rectification types - Estimated nonlinear subunit model of Y-cells explains a significant fraction of response variance to texture movies. - The subunit model also predicts the “Y-cell signature” spatial frequency tuning to grating stimuli. - Estimated Y cell models have varying subunit rectification while X cell models have no rectification. Spatial Frequency Tuning Linear HWR FWR 0 20 40 60 80 Subunit Rectification Type VAF % 0.1 1 2 0.5 1 Actual (F1) Predicted (F1) Spatial Frequency (cpd) Firing Rate (Hz) Spatial Frequency Tuning Demb et al., 2001b Hochstein & Shapley, 1976 “Predicted” responses are measured by running simulations on estimated subunit model of the neuron. 0.1 1 2 4 0.5 0.0 0.5 1.0 1.5 Spatial Frequency (cpd) Firing Rate (Hz) Actual Predicted F1 F2 0.0 ms 13.3 ms 26.7 ms 40.0 ms 53.3 ms ! = 0.125 ° 1° 0.0 ms 13.3 ms 26.7 ms 40.0 ms 53.3 ms 66.7 ms 80.0 ms 93.3 ms ! = 0.25° 1° ! = 0.25 ° VAF = 64% ! = 0.125 ° ! = 0.25 ° VAF = 62% VAF = 49.6% VAF = 37% ! = 0.375 ° 0.5° 3° 2.5° 2.5° 0.1 1 0.5 2 4 0 2 4 6 8 10 Linear (F1) NonLinear (F2) Spatial Frequency (cpd) Firing Rate (Hz)

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Page 1: Amol Gharat and Curtis Bakeramolgharat.com/model nonlinear Y cell receptive field estimation.pdf · Amol Gharat1and Curtis Baker2 METHODS) RESULTS) CONCLUSIONS) 1. Rosenberg et al

INTRODUCTION  

0.1 10.5 2 40

2

4

6

8

10Linear (F1)NonLinear (F2)

Spatial Frequency (cpd)

Firin

g R

ate

(Hz)

Y-cell responses to texture stimuli predicted by a nonlinear receptive field model based on bipolar cell subunits

1Integrated Program in Neuroscience, McGill Univ.; 2Dept of Ophthalmology, McGill Vision Res. Unit, McGill University, Montreal, Canada Amol Gharat1and Curtis Baker2

METHODS  

RESULTS  

CONCLUSIONS   1.  Rosenberg et al (2010). Subcortical representation of non-fourier image features. J Neuroscience. 30(6):1985-1993. 2.  Demb et al (2001a). Cellular basis for the response to second-order motion cues in Y retinal ganglion cells. Neuron. 32:711-721. 3.  Demb et al (2001b). Bipolar cells contribute to nonlinear spatial summation in the brisk-transient (Y) ganglion cell in mammalian retina. J

Neuroscience. 21(19):7447-7454. 4.  Borghuis et al (2013) Two-photon imaging of nonlinear glutamate release dynamics at bipolar cell synapses in the mouse retina. J

Neuroscience. 33(27):10972-10985. 5.  Hochstein & Shapley (1976). Linear and nonlinear spatial subunits in Y cat retinal ganglion cells. 262:265-284. 6.  http://strflab.berkeley.edu/ Funded by CIHR grant MOP-119498 to CB. Vanier Canada Graduate Scholarship to AG. Many thanks to Guangxing Li and Vargha Talebi.

Y cell “subunit” receptive field X cell receptive field

Variety of Y cell receptive fields

time

Y-cell model

Model Spatial Tuning

1 2 4 80.50.2535

40

45

50

55

Subunit Size

VAF

%

Linear HWR FWR0

20

40

60

Subunit Rectification Type

VAF

%

-  A prime aim of visual neuroscience is to estimate neuronal receptive field models that can predict responses to arbitrary stimuli.

-  X (linear) and Y (non-linear) type cells form two major categories in mammalian retina and LGN.

-  Y type cells may be an important early stage for processing second-order information in visual scenes (Demb et al., 2001a; Rosenberg et al., 2010).

-  A linear receptive field model with static output nonlinearity (LN model) cannot account for nonlinear responses of Y-cells to high resolution texture stimuli.

-  A model with spatial pooling of rectified inputs from several small subunit receptive fields (Demb et al., 2001b) might better predict responses of Y-cells to arbitrary texture stimuli.

Stimuli -  Synthetic Naturalistic Texture

images at 75Hz -  3 datasets training (7500 images) regularization (1875 images) validation (1875 images) -  Sinusoidal gratings (Drifting or

Contrast reversing) Animal Preparation -  Anesthetized and paralyzed cats -  Extracellular, single-unit recordings in LGN using single &

multi-channel electrodes -  Manual spike sorting using Plexon Offline Sorter &

SpikeSorter (UBC) Classification of Cells (X or Y) -  Measured responses of neurons to contrast reversing

gratings at series of spatial frequencies -  If response at second harmonic (F2) was greater than first

harmonic (F1), neuron was classified as Y-type Receptive Field Estimation -  Preprocessing with Gaussian subunits -  Estimation of subunit weights with gradient descent

optimization.(STRFlab Software) -  Regularization (early stopping) -  Measure model performance for series of subunit sizes

and rectification types

-  Estimated nonlinear subunit model of Y-cells explains a significant fraction of response variance to texture movies.

-  The subunit model also predicts the “Y-cell signature” spatial frequency tuning to grating stimuli.

-  Estimated Y cell models have varying subunit rectification while X cell models have no rectification.

Spatial Frequency Tuning

Linear HWR FWR0

20

40

60

80

Subunit Rectification Type

VAF

%

0.1 1 20.5

1 Actual (F1)Predicted (F1)

Spatial Frequency (cpd)

Firin

g R

ate

(Hz)

Spatial Frequency Tuning

Demb et al., 2001b

Hochstein & Shapley, 1976

“Predicted” responses are measured by running simulations on estimated subunit model of the neuron.

0.1 1 2 40.50.0

0.5

1.0

1.5

Spatial Frequency (cpd)

Firin

g R

ate

(Hz)

ActualPredicted

F1 F2

0.0 ms 13.3 ms 26.7 ms

40.0 ms 53.3 ms

! = 0.125°

0.0 ms 13.3 ms 26.7 ms

40.0 ms 53.3 ms 66.7 ms

80.0 ms 93.3 ms

! = 0.25°

! = 0.25°

VAF = 64%

! = 0.125° ! = 0.25°

VAF = 62% VAF = 49.6% VAF = 37%

! = 0.375°

0.5°3° 2.5° 2.5°

0.1 10.5 2 40

2

4

6

8

10Linear (F1)NonLinear (F2)

Spatial Frequency (cpd)

Firin

g R

ate

(Hz)