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Learning visual motion Statistical models of spike trains Learning visual motion with recurrent neural networks Marius Pachitariu Gatsby Unit, UCL adviser: Maneesh Sahani Marius Pachitariu Learning visual motion with RNNs 1 / 48

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Page 1: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Learning visual motion with recurrent neural networks

Marius Pachitariu

Gatsby Unit, UCLadviser: Maneesh Sahani

Marius Pachitariu Learning visual motion with RNNs 1 / 48

Page 2: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Outline

Learning visual motionSpatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Statistical models of spike trainsRecurrent GLMInstantaneous noiseResults

Marius Pachitariu Learning visual motion with RNNs 2 / 48

Page 3: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Marr’s three levels of analysis

Levels of analysis

I Computational

I Algorithmic / Representational

I Physical

Marius Pachitariu Learning visual motion with RNNs 3 / 48

Page 4: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Sequential data types

I Movies

I Spike trains

I Language

Marius Pachitariu Learning visual motion with RNNs 4 / 48

Page 5: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Spatio-temporal filters

I Dominant in both visual neuroscience andcomputer vision.

I Caveats:I not real-time/requires copies of the past→ bad for real-world systems, like the brain.

I too many parameters→ bad for learning and generalization.

I high computational complexity→ bad with high-bandwidth data.

Marius Pachitariu Learning visual motion with RNNs 5 / 48

Page 6: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Spatio-temporal filters

I Dominant in both visual neuroscience andcomputer vision.

I Caveats:I not real-time/requires copies of the past→ bad for real-world systems, like the brain.

I too many parameters→ bad for learning and generalization.

I high computational complexity→ bad with high-bandwidth data.

Marius Pachitariu Learning visual motion with RNNs 5 / 48

Page 7: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Neural candidates for ST filters

I lagged LGN cells (Mastronarde, 1987)

Marius Pachitariu Learning visual motion with RNNs 6 / 48

Page 8: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Neural candidates for ST filters

I lagged LGN cells (Mastronarde, 1987)

I but LGN is an information bottleneck

I but LGN responds precisely to natural movies (Butts et al, 2011)

Marius Pachitariu Learning visual motion with RNNs 7 / 48

Page 9: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Neural candidates for ST filters

I lagged LGN cells (Mastronarde, 1987)

I but LGN is an information bottleneck

I but LGN responds precisely to natural movies (Butts et al, 2011)

Marius Pachitariu Learning visual motion with RNNs 7 / 48

Page 10: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Compact parametrization of ST filters with an RNN

xt

ytyt−1yt−2yt−τ ••••••

W0W1W2Wτ

Spatiotemporal filtering

xtxt−1xt−2xt−τ

ytyt−1yt−2yt−τ

••••••

••••••

W0W0W0W0

RRRRR

Recurrent neural network

xt =∞∑τ=0

Wτ yt−τ

Wτ = (R)τ W0

xt = W0 yt + R∞∑τ=0

(R)τ W0 yt−1−τ

xt = W0 yt + R xt−1

As a simple example, we fit Rto a diverse bank ofspatiotemporal filters.

Marius Pachitariu Learning visual motion with RNNs 8 / 48

Page 11: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Compact parametrization of ST filters with an RNN

xt

ytyt−1yt−2yt−τ ••••••

W0W1W2Wτ

Spatiotemporal filtering

xtxt−1xt−2xt−τ

ytyt−1yt−2yt−τ

••••••

••••••

W0W0W0W0

RRRRR

Recurrent neural network

xt =∞∑τ=0

Wτ yt−τ

Wτ = (R)τ W0

xt = W0 yt + R∞∑τ=0

(R)τ W0 yt−1−τ

xt = W0 yt + R xt−1

As a simple example, we fit Rto a diverse bank ofspatiotemporal filters.

Marius Pachitariu Learning visual motion with RNNs 8 / 48

Page 12: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Compact parametrization of ST filters with an RNN

xt

ytyt−1yt−2yt−τ ••••••

W0W1W2Wτ

Spatiotemporal filtering

xtxt−1xt−2xt−τ

ytyt−1yt−2yt−τ

••••••

••••••

W0W0W0W0

RRRRR

Recurrent neural network

xt =∞∑τ=0

Wτ yt−τ

Wτ = (R)τ W0

xt = W0 yt + R∞∑τ=0

(R)τ W0 yt−1−τ

xt = W0 yt + R xt−1

As a simple example, we fit Rto a diverse bank ofspatiotemporal filters.

Marius Pachitariu Learning visual motion with RNNs 8 / 48

Page 13: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Reconstructions of the ST filters are good

Marius Pachitariu Learning visual motion with RNNs 9 / 48

Page 14: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

What do the connections look like?

I Spectrum of R

I Strongest connections to a given neuron (animation).

Marius Pachitariu Learning visual motion with RNNs 10 / 48

Page 15: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

What do the connections look like?

I Spectrum of R

I Strongest connections to a given neuron (animation).

Marius Pachitariu Learning visual motion with RNNs 10 / 48

Page 16: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Computational complexity and memory requirements

I lx by ly by nt filters (12 by 12 by 30)

I N (1600) ST filters

I Feedforward flops = 2N l2x l2

y nt

I Recurrent flops = 2N2 + 2N l2x l2

y

I 5 % non-zero connections in R.

I Recurrent flops = 2 · 0.05 · N2 + 2N l2x l2

y

I Recurrent flops < Feedforward flops

Marius Pachitariu Learning visual motion with RNNs 11 / 48

Page 17: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Computational complexity and memory requirements

I lx by ly by nt filters (12 by 12 by 30)

I N (1600) ST filters

I Feedforward flops = 2N l2x l2

y nt

I Recurrent flops = 2N2 + 2N l2x l2

y

I 5 % non-zero connections in R.

I Recurrent flops = 2 · 0.05 · N2 + 2N l2x l2

y

I Recurrent flops < Feedforward flops

Marius Pachitariu Learning visual motion with RNNs 11 / 48

Page 18: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Computational complexity and memory requirements

I lx by ly by nt filters (12 by 12 by 30)

I N (1600) ST filters

I Feedforward flops = 2N l2x l2

y nt

I Recurrent flops = 2N2 + 2N l2x l2

y

I 5 % non-zero connections in R.

I Recurrent flops = 2 · 0.05 · N2 + 2N l2x l2

y

I Recurrent flops < Feedforward flops

Marius Pachitariu Learning visual motion with RNNs 11 / 48

Page 19: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Computational complexity and memory requirements

I lx by ly by nt filters (12 by 12 by 30)

I N (1600) ST filters

I Feedforward flops = 2N l2x l2

y nt

I Recurrent flops = 2N2 + 2N l2x l2

y

I 5 % non-zero connections in R.

I Recurrent flops = 2 · 0.05 · N2 + 2N l2x l2

y

I Recurrent flops < Feedforward flops

Marius Pachitariu Learning visual motion with RNNs 11 / 48

Page 20: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Advantages of recurrent neural networks

I the brain already has the hardware

I do not require copies of the past→ less memory usage→ the brain has short timescales + bottleneck in LGN→ no evidence for true delay lines in cortex

I fewer parameters→ important for learning and generalization

I reduced computational complexity→ good for high bandwidth data

I can integrate over long time periods→ natural visual motion can be slow and noisy

Marius Pachitariu Learning visual motion with RNNs 12 / 48

Page 21: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Advantages of recurrent neural networks

I the brain already has the hardware

I do not require copies of the past→ less memory usage→ the brain has short timescales + bottleneck in LGN→ no evidence for true delay lines in cortex

I fewer parameters→ important for learning and generalization

I reduced computational complexity→ good for high bandwidth data

I can integrate over long time periods→ natural visual motion can be slow and noisy

Marius Pachitariu Learning visual motion with RNNs 12 / 48

Page 22: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Advantages of recurrent neural networks

I the brain already has the hardware

I do not require copies of the past→ less memory usage→ the brain has short timescales + bottleneck in LGN→ no evidence for true delay lines in cortex

I fewer parameters→ important for learning and generalization

I reduced computational complexity→ good for high bandwidth data

I can integrate over long time periods→ natural visual motion can be slow and noisy

Marius Pachitariu Learning visual motion with RNNs 12 / 48

Page 23: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Advantages of recurrent neural networks

I the brain already has the hardware

I do not require copies of the past→ less memory usage→ the brain has short timescales + bottleneck in LGN→ no evidence for true delay lines in cortex

I fewer parameters→ important for learning and generalization

I reduced computational complexity→ good for high bandwidth data

I can integrate over long time periods→ natural visual motion can be slow and noisy

Marius Pachitariu Learning visual motion with RNNs 12 / 48

Page 24: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Advantages of recurrent neural networks

I the brain already has the hardware

I do not require copies of the past→ less memory usage→ the brain has short timescales + bottleneck in LGN→ no evidence for true delay lines in cortex

I fewer parameters→ important for learning and generalization

I reduced computational complexity→ good for high bandwidth data

I can integrate over long time periods→ natural visual motion can be slow and noisy

Marius Pachitariu Learning visual motion with RNNs 12 / 48

Page 25: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Neural sequence learning via STDP (toy model)

Rao & Sejnowski, NIPS 2000

Marius Pachitariu Learning visual motion with RNNs 13 / 48

Page 26: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Spike and Slab Sparse Coding

I Olshausen&Millman 2000, Rehn&Sommer 2007, Goodfellow&al 2012

zt

xt

ht

yt

pτx

W, τy

htk = Bernoulli (pk)

xt = N(

0, τ 2x · I)

zt = ht ◦ xt

yt = N(W · zt , τ 2

y

)

Marius Pachitariu Learning visual motion with RNNs 14 / 48

Page 27: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Spike and Slab Sparse Coding

I Olshausen&Millman 2000, Rehn&Sommer 2007, Goodfellow&al 2012

zt

xt

ht

yt

pτx

W, τy

htk = Bernoulli (pk)

xt = N(

0, τ 2x · I)

zt = ht ◦ xt

yt = N(W · zt , τ 2

y

)

Marius Pachitariu Learning visual motion with RNNs 14 / 48

Page 28: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Spike and Slab Recurrent Neural Network

P(ht+1|zt

)= σ

(R · zt + b

)σ(x) = 1/(1 + exp(−x))

zt

xt

ht

yt

ht+1

zt−1

t

R,bτx

W, τy

. . .. . .

For approximate inference:

I Assuming we have setxt , ht for t = 1 to T .

I At T + 1 we only needto solve an SCproblem.

Marius Pachitariu Learning visual motion with RNNs 15 / 48

Page 29: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Spike and Slab Recurrent Neural Network

P(ht+1|zt

)= σ

(R · zt + b

)σ(x) = 1/(1 + exp(−x))

zt

xt

ht

yt

ht+1

zt−1

t

R,bτx

W, τy

. . .. . .

For approximate inference:

I Assuming we have setxt , ht for t = 1 to T .

I At T + 1 we only needto solve an SCproblem.

Marius Pachitariu Learning visual motion with RNNs 15 / 48

Page 30: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Natural movies and artificial stimuli

Training data (ZCA whitened) Full single frame of training data

Test data (whitened)

Marius Pachitariu Learning visual motion with RNNs 16 / 48

Page 31: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Results - Speed tuning

Speed (pix/frame)

Response(a.u.)

preferred non-preferred

0 3

Rectified neural responses max(z, 0) todrifting square gratings.

Orban et al, 1986

Marius Pachitariu Learning visual motion with RNNs 17 / 48

Page 32: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Results - Speed tuning

Speed (pix/frame)

Response(a.u.)

preferred non-preferred

0 3

Rectified neural responses max(z, 0) todrifting square gratings.

Orban et al, 1986

Marius Pachitariu Learning visual motion with RNNs 17 / 48

Page 33: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Results - Direction selectivity indices

0 1 2 30

50

100

150

Numberofunits

Preferred speed (pix/frame)

Direction Index0 0.5 1

0

100

200

DI = 1− rnon-pref/rpref

Peterson et al, 2004

Gur et al, 2007

Marius Pachitariu Learning visual motion with RNNs 18 / 48

Page 34: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Results - Direction selectivity indices

0 1 2 30

50

100

150

Numberofunits

Preferred speed (pix/frame)

Direction Index0 0.5 1

0

100

200

DI = 1− rnon-pref/rpref

Peterson et al, 2004

Gur et al, 2007

Marius Pachitariu Learning visual motion with RNNs 18 / 48

Page 35: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Results - Direction selectivity indices

0 1 2 30

50

100

150

Numberofunits

Preferred speed (pix/frame)

Direction Index0 0.5 1

0

100

200

DI = 1− rnon-pref/rpref

Peterson et al, 2004

Gur et al, 2007

Marius Pachitariu Learning visual motion with RNNs 18 / 48

Page 36: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Results - connectomics in silico

Largest outgoing connections of oneunit

Presynaptic

Postsynaptic

0

0

Connected units are co-oriented

Marius Pachitariu Learning visual motion with RNNs 19 / 48

Page 37: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Results - connectomics in silico

Largest outgoing connections of oneunit

Presynaptic

Postsynaptic

0

0

Connected units are co-oriented

Marius Pachitariu Learning visual motion with RNNs 19 / 48

Page 38: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Results - connectomics in silico

Outgoing connections of 15randomly chosen DS units (andanimation during learning)

Responses to small drifting Gabors- polar plots

Marius Pachitariu Learning visual motion with RNNs 20 / 48

Page 39: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Results - connectomics in silico

Outgoing connections of 15randomly chosen DS units (andanimation during learning)

Responses to small drifting Gabors- polar plots

Marius Pachitariu Learning visual motion with RNNs 20 / 48

Page 40: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Inference and learning

zt

x tht

y t

ht+ 1

zt− 1

t

R , bτx

W , τy

. . .. . .

Lss-RNN =∑t

Ltss-RNN

Ltss-RNN = const− ‖yt −W(xt ◦ ht)‖2/2τ 2

y − ‖xt‖2/2τ 2x +

+N∑j=1

htj log σ

(R(ht−1 ◦ xt−1

)+ b)j

+N∑j=1

(1− htj ) log

(1− σ

(R(ht−1 ◦ xt−1

)+ b)j

)

For approximate inference, we use greedy filtering:

I Assuming we have already set xt , ht for t = 1 to T .

I At step T + 1 we only need to solve an sparse codingproblem given by the slice LT+1

ss-RNN.

I We solve the SC problem with standard matchingpursuit / coordinate descent methods.

Marius Pachitariu Learning visual motion with RNNs 21 / 48

Page 41: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Inference and learning

zt

x tht

y t

ht+ 1

zt− 1

t

R , bτx

W , τy

. . .. . .

Lss-RNN =∑t

Ltss-RNN

Ltss-RNN = const− ‖yt −W(xt ◦ ht)‖2/2τ 2

y − ‖xt‖2/2τ 2x +

+N∑j=1

htj log σ

(R(ht−1 ◦ xt−1

)+ b)j

+N∑j=1

(1− htj ) log

(1− σ

(R(ht−1 ◦ xt−1

)+ b)j

)For approximate inference, we use greedy filtering:

I Assuming we have already set xt , ht for t = 1 to T .

I At step T + 1 we only need to solve an sparse codingproblem given by the slice LT+1

ss-RNN.

I We solve the SC problem with standard matchingpursuit / coordinate descent methods.

Marius Pachitariu Learning visual motion with RNNs 21 / 48

Page 42: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Learning of ss-RNN

Rao & Sejnowski, NIPS 2000

Gradients for learning R are similar to the STDP learning rule used in Rao &Sejnowski, 2000.

∂Ltss-RNN

∂Rjk=(

ht−1k x t−1

k

)·(

htj − σ

(R(ht ◦ xt

)+ b)j

).

Marius Pachitariu Learning visual motion with RNNs 22 / 48

Page 43: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

DS is learned, OS is not?

I Orientation selectivity (OS) and ocular dominance (OD) do not requirevisual experience

I Visual deprivation has little impact on OS and OD

I However, DS does require visual experience in ferrets (Li et al, 2006)

Marius Pachitariu Learning visual motion with RNNs 23 / 48

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Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

DS is learned, OS is not?

I Orientation selectivity (OS) and ocular dominance (OD) do not requirevisual experience

I Visual deprivation has little impact on OS and OD

I However, DS does require visual experience in ferrets (Li et al, 2006)

Marius Pachitariu Learning visual motion with RNNs 23 / 48

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Learning visual motionStatistical models of spike trains

Spatiotemporal filteringRecurrent neural networks can compute visual motionLearning in generative RNN

Conclusions

I Recurrent neural networks can analyze visual motion in an online fashionwithout delayed inputs.

I Formulating a generative model allows learning the recurrent connectionsvia an STDP rule.

I As a model of V1, the RNN makes testable predictions about the lateralconnectivity of neurons.

I Responses to stimuli may however be similar to those of spatiotemporalfilters.

Marius Pachitariu Learning visual motion with RNNs 24 / 48

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Learning visual motionStatistical models of spike trains

Recurrent GLMInstantaneous noiseResults

Statement of the problem

Marius Pachitariu Learning visual motion with RNNs 25 / 48

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Learning visual motionStatistical models of spike trains

Recurrent GLMInstantaneous noiseResults

The equivalent of spatiotemporal filters:Generalized Linear Models (GLMs)

Pillow et al, 2008.

Marius Pachitariu Learning visual motion with RNNs 26 / 48

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Learning visual motionStatistical models of spike trains

Recurrent GLMInstantaneous noiseResults

Recurrent Generalized Linear Models

xtxt−1xt−2xt−τ

ytyt−1yt−2yt−τ

••••••

••••••

W0W0W0W0

RRRRR

ZZ

predictive RNN

xt = W0 yt + R xt−1

yt ⊥⊥ {yt−1, yt−2, ...} | xt

Similar to state-of-the-art language models: Sutskever et al, 2011, Mikolov etal, 2011

Marius Pachitariu Learning visual motion with RNNs 27 / 48

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Learning visual motionStatistical models of spike trains

Recurrent GLMInstantaneous noiseResults

Recurrent Generalized Linear Models

xtxt−1xt−2xt−τ

ytyt−1yt−2yt−τ

••••••

••••••

W0W0W0W0

RRRRR

ZZ

predictive RNN

xt = W0 yt + R xt−1

yt ⊥⊥ {yt−1, yt−2, ...} | xt

Similar to state-of-the-art language models: Sutskever et al, 2011, Mikolov etal, 2011

Marius Pachitariu Learning visual motion with RNNs 27 / 48

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Learning visual motionStatistical models of spike trains

Recurrent GLMInstantaneous noiseResults

Relationship to linear dynamical system (LDS)

xtxt−1xt−2xt−τ

ytyt−1yt−2yt−τ

••••••

••••••

W0W0W0W0

RRRRR

ZZ

R-GLM

xtxt−1xt−2xt−τ

ytyt−1yt−2yt−τ

••••••

••••••

W0W0W0W0

RRRRR

LDS

Marius Pachitariu Learning visual motion with RNNs 28 / 48

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Learning visual motionStatistical models of spike trains

Recurrent GLMInstantaneous noiseResults

Problem: cannot have instantaneous connections in a GLM

I where are instantaneous correlations coming from?

I can add Ising observation model butI partition function

p(x) =1

Ze−xTAx/2−xT b

I not available for Poisson observationsI cannot add nonlinear link function like in GLM

Marius Pachitariu Learning visual motion with RNNs 29 / 48

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Learning visual motionStatistical models of spike trains

Recurrent GLMInstantaneous noiseResults

Problem: cannot have instantaneous connections in a GLM

I where are instantaneous correlations coming from?I can add Ising observation model but

I partition function

p(x) =1

Ze−xTAx/2−xT b

I not available for Poisson observationsI cannot add nonlinear link function like in GLM

Marius Pachitariu Learning visual motion with RNNs 29 / 48

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Learning visual motionStatistical models of spike trains

Recurrent GLMInstantaneous noiseResults

Our solution: sequential prediction of each neuron

I constrain A to be strictly lower triangular

p(x) = Poisson (f (Ax + b))

I Can do the same with Gaussian observation noise.→ Equivalent to full covariance Gaussians.

I What about the ordering?

I Can do the same with Bernoulli observation noise.→ Performance matches Ising model.

I Similar to recent image models: Theis et al, 2011 and Larochelle&Murray,2011.

Marius Pachitariu Learning visual motion with RNNs 30 / 48

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Learning visual motionStatistical models of spike trains

Recurrent GLMInstantaneous noiseResults

Our solution: sequential prediction of each neuron

I constrain A to be strictly lower triangular

p(x) = Poisson (f (Ax + b))

I Can do the same with Gaussian observation noise.→ Equivalent to full covariance Gaussians.

I What about the ordering?

I Can do the same with Bernoulli observation noise.→ Performance matches Ising model.

I Similar to recent image models: Theis et al, 2011 and Larochelle&Murray,2011.

Marius Pachitariu Learning visual motion with RNNs 30 / 48

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Learning visual motionStatistical models of spike trains

Recurrent GLMInstantaneous noiseResults

Our solution: sequential prediction of each neuron

I constrain A to be strictly lower triangular

p(x) = Poisson (f (Ax + b))

I Can do the same with Gaussian observation noise.→ Equivalent to full covariance Gaussians.

I What about the ordering?

I Can do the same with Bernoulli observation noise.→ Performance matches Ising model.

I Similar to recent image models: Theis et al, 2011 and Larochelle&Murray,2011.

Marius Pachitariu Learning visual motion with RNNs 30 / 48

Page 56: Learning visual motion with recurrent neural networksmarius/slides/RedwoodTalk.pdf · Marius Pachitariu Learning visual motion with RNNs 12/48. Learning visual motion Statistical

Learning visual motionStatistical models of spike trains

Recurrent GLMInstantaneous noiseResults

Our solution: sequential prediction of each neuron

I constrain A to be strictly lower triangular

p(x) = Poisson (f (Ax + b))

I Can do the same with Gaussian observation noise.→ Equivalent to full covariance Gaussians.

I What about the ordering?

I Can do the same with Bernoulli observation noise.→ Performance matches Ising model.

I Similar to recent image models: Theis et al, 2011 and Larochelle&Murray,2011.

Marius Pachitariu Learning visual motion with RNNs 30 / 48

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Learning visual motionStatistical models of spike trains

Recurrent GLMInstantaneous noiseResults

Our solution: sequential prediction of each neuron

I constrain A to be strictly lower triangular

p(x) = Poisson (f (Ax + b))

I Can do the same with Gaussian observation noise.→ Equivalent to full covariance Gaussians.

I What about the ordering?

I Can do the same with Bernoulli observation noise.→ Performance matches Ising model.

I Similar to recent image models: Theis et al, 2011 and Larochelle&Murray,2011.

Marius Pachitariu Learning visual motion with RNNs 30 / 48

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Learning visual motionStatistical models of spike trains

Recurrent GLMInstantaneous noiseResults

Sampling the correlated Poisson model

Data correlations Model correlations

Marius Pachitariu Learning visual motion with RNNs 31 / 48

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Learning visual motionStatistical models of spike trains

Recurrent GLMInstantaneous noiseResults

Instantaneous noise and recurrence increase the likelihood

Likelihood(bits/spike)

fully independent - 3.15correlated Poisson + 0.175GLM + 0.225GLM with correlated Pois-son

+ 0.03

R-GLM with correlatedPoisson

+ 0.03

Marius Pachitariu Learning visual motion with RNNs 32 / 48

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Learning visual motionStatistical models of spike trains

Recurrent GLMInstantaneous noiseResults

R-GLM learns long timescales

0.85 0.9 0.95 1−0.4

−0.2

0

0.2

0.4

real

imag

inar

y

R−GLMLDS

Eigenvalues of the recurrent matrix

Marius Pachitariu Learning visual motion with RNNs 33 / 48

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Learning visual motionStatistical models of spike trains

Recurrent GLMInstantaneous noiseResults

Samples of model reproduce spatiotemporal correlations in data

LDS and GLM

−100 −50 0 50 100−0.01

0

0.01

0.02

0.03

0.04

0.05

0.06

Time lag (ms)

Cor

rela

tion

Neurons 1−23Neurons 24−46Neurons 47−69Neurons 70−92

R-GLM

Marius Pachitariu Learning visual motion with RNNs 34 / 48

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Learning visual motionStatistical models of spike trains

Recurrent GLMInstantaneous noiseResults

Samples of model reproduce spatiotemporal correlations in data

LDS and GLM

−100 −50 0 50 100−0.01

0

0.01

0.02

0.03

0.04

0.05

0.06

Time lag (ms)

Cor

rela

tion

Neurons 1−23Neurons 24−46Neurons 47−69Neurons 70−92

R-GLM

Marius Pachitariu Learning visual motion with RNNs 34 / 48

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Learning visual motionStatistical models of spike trains

Recurrent GLMInstantaneous noiseResults

Hidden units integrate info about stimulus

0 200 400 600 800 1000 1200

Simulation

Time (ms)

0 200 400 600 800 1000 1200

Data

Time (ms)

Delay period

Marius Pachitariu Learning visual motion with RNNs 35 / 48

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Learning visual motionStatistical models of spike trains

Recurrent GLMInstantaneous noiseResults

Hidden units generate dynamics

0 200 400 600 800 1000 1200 1400 1600 1800

Simulation

Time (ms)

0 200 400 600 800 1000 1200 1400 1600 1800

Data

Time (ms)

Delay + Movement period

Marius Pachitariu Learning visual motion with RNNs 36 / 48

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Learning visual motionStatistical models of spike trains

Recurrent GLMInstantaneous noiseResults

Joint estimation with hand position improves decoding

xtxt−1xt−2xt−τ

ytyt−1yt−2yt−τ

••••••

••••••

W0W0W0W0

RRRRR

ZZ

Task

Mixture of trajectories model

Our result: 6.45 mm.

Marius Pachitariu Learning visual motion with RNNs 37 / 48

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Learning visual motionStatistical models of spike trains

Recurrent GLMInstantaneous noiseResults

Joint estimation with hand position improves decoding

xtxt−1xt−2xt−τ

ytyt−1yt−2yt−τ

••••••

••••••

W0W0W0W0

RRRRR

ZZ

Task Mixture of trajectories model

Our result: 6.45 mm.

Marius Pachitariu Learning visual motion with RNNs 37 / 48

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Learning visual motionStatistical models of spike trains

Recurrent GLMInstantaneous noiseResults

Joint estimation with hand position improves decoding

xtxt−1xt−2xt−τ

ytyt−1yt−2yt−τ

••••••

••••••

W0W0W0W0

RRRRR

ZZ

Task Mixture of trajectories model

Our result: 6.45 mm.

Marius Pachitariu Learning visual motion with RNNs 37 / 48

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Learning visual motionStatistical models of spike trains

Recurrent GLMInstantaneous noiseResults

Speed profiles

0 10 20 300

5

10

15

0 10 20 300

5

10

15

0 10 20 300

5

10

15

0 10 200

5

10

15

0 10 20 300

5

10

15

0 10 20 300

5

10

15

0 10 20 300

5

10

15

0 10 200

5

10

15

0 10 20 300

5

10

15

0 10 20 300

5

10

15

0 10 20 300

5

10

15

0 10 200

5

10

15

0 10 20 300

5

10

15

0 10 20 300

5

10

15

0 10 20 300

5

10

15

0 10 200

5

10

15

Marius Pachitariu Learning visual motion with RNNs 38 / 48

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Learning visual motionStatistical models of spike trains

Recurrent GLMInstantaneous noiseResults

Conclusions

I Recurrent GLMs with correlated Poisson observations improve statisticalmodels of spike trains.

I The low dimensional parametrization improves decoding of handtrajectories from neural data.

I This work was funded by the Gatsby Charitable Foundation.

Marius Pachitariu Learning visual motion with RNNs 39 / 48

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Learning visual motionStatistical models of spike trains

Recurrent GLMInstantaneous noiseResults

The Bayesian Sampling hypothesis

I Bayesian brain hypothesis

I how does the brain do inference: sampling

I interesting data from V1 supports Bayesian sampling hypothesis

I visual word reading time ∼ − log(P(word))

I Bayesian reader (Norris, 2006) collects visual samples untilP(word|visual samples) is large.

Marius Pachitariu Learning visual motion with RNNs 40 / 48

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Learning visual motionStatistical models of spike trains

Recurrent GLMInstantaneous noiseResults

The Bayesian Sampling hypothesis

I Bayesian brain hypothesis

I how does the brain do inference: sampling

I interesting data from V1 supports Bayesian sampling hypothesis

I visual word reading time ∼ − log(P(word))

I Bayesian reader (Norris, 2006) collects visual samples untilP(word|visual samples) is large.

Marius Pachitariu Learning visual motion with RNNs 40 / 48

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Learning visual motionStatistical models of spike trains

Recurrent GLMInstantaneous noiseResults

The Bayesian Sampling hypothesis

I Bayesian brain hypothesis

I how does the brain do inference: sampling

I interesting data from V1 supports Bayesian sampling hypothesis

I visual word reading time ∼ − log(P(word))

I Bayesian reader (Norris, 2006) collects visual samples untilP(word|visual samples) is large.

Marius Pachitariu Learning visual motion with RNNs 40 / 48

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Learning visual motionStatistical models of spike trains

Recurrent GLMInstantaneous noiseResults

The sequential Bayesian Reader

I Hypothesis: visual word reading time ∼ P(word|history)I Need two ingredients

I DataI good language models

Marius Pachitariu Learning visual motion with RNNs 41 / 48

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Learning visual motionStatistical models of spike trains

Recurrent GLMInstantaneous noiseResults

Language modelling: statement of the problem

The Great Gatsby, by F. Scott FitzgeraldIn my younger and more vulnerable years my father gave me some advice thatI’ve been turning over in my mind ever since.”Whenever you feel like criticizing any one,” he told me, ”just remember thatall the people in this world haven’t had the advantages that you’ve had.”

Marius Pachitariu Learning visual motion with RNNs 42 / 48

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Learning visual motionStatistical models of spike trains

Recurrent GLMInstantaneous noiseResults

The equivalent of spatio-temporal filters and GLMs: N-grams

4-gram (frequency)

I serve as the incoming (92)

I serve as the incubator (99)

I serve as the independent (794)

I serve as the index (223)

I serve as the indication (72)

I serve as the indicator (120)

Marius Pachitariu Learning visual motion with RNNs 43 / 48

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Learning visual motionStatistical models of spike trains

Recurrent GLMInstantaneous noiseResults

Neural network language models

4-gram (frequency)

I serve as the incoming (92)

I serve as the incubator (99)

I serve as the independent (794)

I serve as the index (223)

I serve as the indication (72)

I serve as the indicator (120)

serve as the incoming

Neural Network Language Model

ht0 = f

(t=∞∑t=1

WT It0−t

)P(It0) = softmax(ZTht0).

Marius Pachitariu Learning visual motion with RNNs 44 / 48

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Learning visual motionStatistical models of spike trains

Recurrent GLMInstantaneous noiseResults

Recurrent neural network language models

serve as the incoming

Neural Network Language Model

serve as the incoming

Recurrent Neural Network LanguageModel

I State of the art (Mikolov et al2011).

I Our simplification: linear RNN(R-GLM).

ht0 = R ht0−1 + WT0 It0−t.

ht0 =t=∞∑t=0

Rt It0−t.

Marius Pachitariu Learning visual motion with RNNs 45 / 48

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Learning visual motionStatistical models of spike trains

Recurrent GLMInstantaneous noiseResults

What are the relevant time scales of language?

I R-GLM learns caching.

0 128 256 384 512

0.25

1

3

10

3060

Hidden dimension #

Tim

e sc

ale

(wor

ds in

the

past

)

The timescales of language

I Long time scales are good: dynamic R-GLM further adapts parameters attest time.

Marius Pachitariu Learning visual motion with RNNs 46 / 48

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Learning visual motionStatistical models of spike trains

Recurrent GLMInstantaneous noiseResults

Perplexity results on Penn Corpus (930k tokens, 10k vocab) - single models

Single +KN5+cache x10 x10+KN5+cache

5-gram Kneser-Ney1 141.2 125.7

feedforward NNLM1 140.2 106.6

Log-bilinear LM1 144.5 105.8

RNN1 124.7 97.5 102.1 89.4

dynamic RNN1 123.2 98.0 101.0 90.0

R-GLM(no reg) 137R-GLM(L1 reg) 125

R-GLM (2DO&CN) 102 94 98.8 92.5

dynamic R-GLM(2DO&CN) 98.4 90.7 95.1 89.1

1 copied from Tomas Mikolov thesis2 trained with random dropouts and column normalization

Marius Pachitariu Learning visual motion with RNNs 47 / 48

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Learning visual motionStatistical models of spike trains

Recurrent GLMInstantaneous noiseResults

Conclusions

I None yet. Need to collect data.

Marius Pachitariu Learning visual motion with RNNs 48 / 48