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A plausible model of recognition and postdiction in dynamic environment

Kevin Li, Maneesh Sahani

Gatsby Computational Neuroscience Unit, University College London

January 6, 2020

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 1 / 20

Introduction

1. Introduction

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 2 / 20

Introduction

Inference using an internal model (Helmholtz machine)

static world

p(z,x) −→ r(·)

z

x

r

xDayan, Hinton, Neal & Zemel, 1995

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 3 / 20

Introduction

Inference using an internal model (Helmholtz machine)

static world

p(z,x) −→ r(·)

z

x

r

x

dynamic world

p(z1:t,x1:t) −→ rt(·)

zt−1 zt

xtxt−1

rt−1 rt

xtxt−1Dayan, Hinton, Neal & Zemel, 1995

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 3 / 20

Introduction

Illusion 1

0 1 2 3time (sec)

500

1000

1500

frequ

ency

(Hz)

0 1 2 3time (sec)

500

1000

1500

frequ

ency

(Hz)

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 4 / 20

Introduction

Illusion 1

0 1 2 3time (sec)

500

1000

1500

frequ

ency

(Hz)

0 1 2 3time (sec)

500

1000

1500

frequ

ency

(Hz)

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 4 / 20

Introduction

Illusion 1

0 1 2 3time (sec)

500

1000

1500

frequ

ency

(Hz)

0 1 2 3time (sec)

500

1000

1500

frequ

ency

(Hz)

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 4 / 20

Introduction

Illusion 1

0 1 2 3time (sec)

500

1000

1500

frequ

ency

(Hz)

0 1 2 3time (sec)

500

1000

1500

frequ

ency

(Hz)

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 4 / 20

Introduction

Illusion 2: cutaneous rabbit

In the course of designing some experiments on thecutaneous perception of mechanical pulses deliveredto the back of the forearm, it was discovered that,under some conditions of timing, the taps producedseemed not to be properly localized under thecontactors. [...] They will seem to be distributed,with more or less uniform spacing, from the regionof the �rst contactor to that of the third. There is

a smooth progression of jumps up the arm, as

if a tiny rabbit were hopping from elbow to wrist.

Geldard & Sherrick, 1972, Science

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 5 / 20

Introduction

Illusion 2: cutaneous rabbit

In the course of designing some experiments on thecutaneous perception of mechanical pulses deliveredto the back of the forearm, it was discovered that,under some conditions of timing, the taps producedseemed not to be properly localized under thecontactors. [...] They will seem to be distributed,with more or less uniform spacing, from the regionof the �rst contactor to that of the third. There is

a smooth progression of jumps up the arm, as

if a tiny rabbit were hopping from elbow to wrist.

Geldard & Sherrick, 1972, Science

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 5 / 20

Introduction

In those examples:

Percept of the past is changed by new, future observation

�Law of Continuity�

What could be the neural basis for these statistical computation?

representing beliefs as distributional objects

updating beliefs of the past based on new evidence in real time?

learning to do all the above

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 6 / 20

Introduction

In those examples:

Percept of the past is changed by new, future observation

�Law of Continuity�

What could be the neural basis for these statistical computation?

representing beliefs as distributional objects

updating beliefs of the past based on new evidence in real time?

learning to do all the above

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 6 / 20

Introduction

In those examples:

Percept of the past is changed by new, future observation

�Law of Continuity�

What could be the neural basis for these statistical computation?

representing beliefs as distributional objects

updating beliefs of the past based on new evidence in real time?

learning to do all the above

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 6 / 20

Introduction

In those examples:

Percept of the past is changed by new, future observation

�Law of Continuity�

What could be the neural basis for these statistical computation?

representing beliefs as distributional objects

updating beliefs of the past based on new evidence in real time?

learning to do all the above

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 6 / 20

Introduction

In those examples:

Percept of the past is changed by new, future observation

�Law of Continuity�

What could be the neural basis for these statistical computation?

representing beliefs as distributional objects

updating beliefs of the past based on new evidence in real time?

learning to do all the above

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 6 / 20

Introduction

In those examples:

Percept of the past is changed by new, future observation

�Law of Continuity�

What could be the neural basis for these statistical computation?

representing beliefs as distributional objects

updating beliefs of the past based on new evidence in real time?

learning to do all the above

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 6 / 20

Introduction

In those examples:

Percept of the past is changed by new, future observation

�Law of Continuity�

What could be the neural basis for these statistical computation?

representing beliefs as distributional objects

updating beliefs of the past based on new evidence in real time?

learning to do all the above

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 6 / 20

Distributed distributional code

2. Distributed distributional code

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 7 / 20

Distributed distributional code

DDC: a framework for neural representation of uncertainty

A DDC encodes a probability distribution:

q(z)

by a set of tuning functions

γ(z) := [γ1 (z) , γ2 (z) , γ3 (z) , ..., γK (z)]

into a set of expectations

r := Eq(z) [γ1 (z) , γ2 (z) , ..., γK (z)]

Zemel, Dayan & Pouget (1998); Sahani & Dayan (2003),Vértes & Sahani (2018)

, , , , , , , ,1 2 ... K[ ], , , , , , , ,

1 2 ... K[ ], , , , , , , ,

1 2 ... K[ ]

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 8 / 20

Distributed distributional code

DDC: a framework for neural representation of uncertainty

A DDC encodes a probability distribution:

q(z)

by a set of tuning functions

γ(z) := [γ1 (z) , γ2 (z) , γ3 (z) , ..., γK (z)]

into a set of expectations

r := Eq(z) [γ1 (z) , γ2 (z) , ..., γK (z)]

Zemel, Dayan & Pouget (1998); Sahani & Dayan (2003),Vértes & Sahani (2018)

, , , , , , , ,1 2 ... K[ ]

, , , , , , , ,1 2 ... K[ ], , , , , , , ,

1 2 ... K[ ]

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 8 / 20

Distributed distributional code

DDC: a framework for neural representation of uncertainty

A DDC encodes a probability distribution:

q(z)

by a set of tuning functions

γ(z) := [γ1 (z) , γ2 (z) , γ3 (z) , ..., γK (z)]

into a set of expectations

r := Eq(z) [γ1 (z) , γ2 (z) , ..., γK (z)]

Zemel, Dayan & Pouget (1998); Sahani & Dayan (2003),Vértes & Sahani (2018)

, , , , , , , ,1 2 ... K[ ], , , , , , , ,

1 2 ... K[ ]

, , , , , , , ,1 2 ... K[ ]

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 8 / 20

Distributed distributional code

DDC: a framework for neural representation of uncertainty

A DDC encodes a probability distribution:

q(z)

by a set of tuning functions

γ(z) := [γ1 (z) , γ2 (z) , γ3 (z) , ..., γK (z)]

into a set of expectations

r := Eq(z) [γ1 (z) , γ2 (z) , ..., γK (z)]

Zemel, Dayan & Pouget (1998); Sahani & Dayan (2003),Vértes & Sahani (2018)

, , , , , , , ,1 2 ... K[ ], , , , , , , ,

1 2 ... K[ ], , , , , , , ,

1 2 ... K[ ]

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 8 / 20

Distributed distributional code

DDC: a framework for neural representation of uncertainty

A DDC encodes a probability distribution:

q(z)

by a set of tuning functions

γ(z) := [γ1 (z) , γ2 (z) , γ3 (z) , ..., γK (z)]

into a set of expectations

r := Eq(z) [γ1 (z) , γ2 (z) , ..., γK (z)]

Zemel, Dayan & Pouget (1998); Sahani & Dayan (2003),Vértes & Sahani (2018)

, , , , , , , ,1 2 ... K[ ], , , , , , , ,

1 2 ... K[ ], , , , , , , ,

1 2 ... K[ ]

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 8 / 20

Distributed distributional code

Why DDC? It can encode a large family of distributions

given Eq [γk (z)] = rk, ∀k ∈ {1, 2, . . .K}

q(z) = arg maxH[q]

=1

Z(r)exp

[∑k

θk(r)γk(z)

]

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 9 / 20

Distributed distributional code

Why DDC? It can encode a large family of distributions

given Eq [γk (z)] = rk, ∀k ∈ {1, 2, . . .K}q(z) = arg maxH[q]

=1

Z(r)exp

[∑k

θk(r)γk(z)

]

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 9 / 20

Distributed distributional code

Why DDC? It can encode a large family of distributions

given Eq [γk (z)] = rk, ∀k ∈ {1, 2, . . .K}q(z) = arg maxH[q]

=1

Z(r)exp

[∑k

θk(r)γk(z)

]

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 9 / 20

Distributed distributional code

Why DDC? It can encode a large family of distributions

given Eq [γk (z)] = rk, ∀k ∈ {1, 2, . . .K}q(z) = arg maxH[q]

=1

Z(r)exp

[∑k

θk(r)γk(z)

]Normal q(z)

(z)

,[ ]rk

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 9 / 20

Distributed distributional code

Why DDC? It can encode a large family of distributions

given Eq [γk (z)] = rk, ∀k ∈ {1, 2, . . .K}q(z) = arg maxH[q]

=1

Z(r)exp

[∑k

θk(r)γk(z)

]

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 9 / 20

Distributed distributional code

Why DDC? It can encode a large family of distributions

given Eq [γk (z)] = rk, ∀k ∈ {1, 2, . . .K}q(z) = arg maxH[q]

=1

Z(r)exp

[∑k

θk(r)γk(z)

]Discrete q(z)

(z)

, , , , , , , , ,[ ]rk

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 9 / 20

Distributed distributional code

Why DDC? It can encode a large family of distributions

given Eq [γk (z)] = rk, ∀k ∈ {1, 2, . . .K}q(z) = arg maxH[q]

=1

Z(r)exp

[∑k

θk(r)γk(z)

]arbitrary q(z)

q(z)

k(z)

, , , , , , ,[ ]rk

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 9 / 20

Distributed distributional code

Why DDC? It can encode a large family of distributions

given Eq [γk (z)] = rk, ∀k ∈ {1, 2, . . .K}q(z) = arg maxH[q]

=1

Z(r)exp

[∑k

θk(r)γk(z)

]arbitrary q(z)

q(z)q(z)

k(z), K=8

, , , , , , ,[ ]rk

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 9 / 20

Distributed distributional code

Why DDC? It can encode a large family of distributions

given Eq [γk (z)] = rk, ∀k ∈ {1, 2, . . .K}q(z) = arg maxH[q]

=1

Z(r)exp

[∑k

θk(r)γk(z)

]arbitrary q(z)

q(z)q(z)

k(z), K=12

, , , , , , , , , , ,[ ]rk

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 9 / 20

Distributed distributional code

Why DDC? It can encode a large family of distributions

given Eq [γk (z)] = rk, ∀k ∈ {1, 2, . . .K}q(z) = arg maxH[q]

=1

Z(r)exp

[∑k

θk(r)γk(z)

]arbitrary q(z)

q(z)q(z)

k(z), K=16

, , , , , , , , , , , , , , ,[ ]rk

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 9 / 20

Distributed distributional code

Why DDC? It makes computation simple for neurons

r := Eq(z) [γ1 (z) , γ2 (z) , ..., γK (z)]

r := Eq(z) [γ1 (z) , γ2 (z) , ..., γK (z)]

Key computations involve expected values:

Message passing:q(z2) = Eq(z1) [p(z2|z1)]

Marginalization:

q(z2 ∈ (a, b)) = Eq(z1,z2) [1(a < z2 < b)]

Action evaluation:Q(a) = Eq(s) [R(s, a)]

How to compute E [h(z)]?

if h(z) ≈∑i

αiγi(z)⇒ E [h(z)] ≈∑i

αiri

, , , , , , , ,1 2 ... K[ ]

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 10 / 20

Distributed distributional code

Why DDC? It makes computation simple for neurons

r := Eq(z) [γ1 (z) , γ2 (z) , ..., γK (z)]

Key computations involve expected values:

Message passing:q(z2) = Eq(z1) [p(z2|z1)]

Marginalization:

q(z2 ∈ (a, b)) = Eq(z1,z2) [1(a < z2 < b)]

Action evaluation:Q(a) = Eq(s) [R(s, a)]

How to compute E [h(z)]?

if h(z) ≈∑i

αiγi(z)⇒ E [h(z)] ≈∑i

αiri

q(z)

(z)

, , , , , , , ,1 2 ... K[ ]r

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 10 / 20

Distributed distributional code

Why DDC? It makes computation simple for neurons

r := Eq(z) [γ1 (z) , γ2 (z) , ..., γK (z)]

Key computations involve expected values:

Message passing:q(z2) = Eq(z1) [p(z2|z1)]

Marginalization:

q(z2 ∈ (a, b)) = Eq(z1,z2) [1(a < z2 < b)]

Action evaluation:Q(a) = Eq(s) [R(s, a)]

How to compute E [h(z)]?

if h(z) ≈∑i

αiγi(z)⇒ E [h(z)] ≈∑i

αiri

q(z)

(z)

, , , , , , , ,1 2 ... K[ ]r

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 10 / 20

Distributed distributional code

Why DDC? It makes computation simple for neurons

r := Eq(z) [γ1 (z) , γ2 (z) , ..., γK (z)]

Key computations involve expected values:

Message passing:q(z2) = Eq(z1) [p(z2|z1)]

Marginalization:

q(z2 ∈ (a, b)) = Eq(z1,z2) [1(a < z2 < b)]

Action evaluation:Q(a) = Eq(s) [R(s, a)]

How to compute E [h(z)]?

if h(z) ≈∑i

αiγi(z)⇒ E [h(z)] ≈∑i

αiri

q(z)

(z)

, , , , , , , ,1 2 ... K[ ]r

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 10 / 20

Distributed distributional code

Why DDC? It makes computation simple for neurons

r := Eq(z) [γ1 (z) , γ2 (z) , ..., γK (z)]

Key computations involve expected values:

Message passing:q(z2) = Eq(z1) [p(z2|z1)]

Marginalization:

q(z2 ∈ (a, b)) = Eq(z1,z2) [1(a < z2 < b)]

Action evaluation:Q(a) = Eq(s) [R(s, a)]

How to compute E [h(z)]?

if h(z) ≈∑i

αiγi(z)⇒ E [h(z)] ≈∑i

αiri

q(z)

(z)

, , , , , , , ,1 2 ... K[ ]r

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 10 / 20

Distributed distributional code

Why DDC? It makes computation simple for neurons

r := Eq(z) [γ1 (z) , γ2 (z) , ..., γK (z)]

Key computations involve expected values:

Message passing:q(z2) = Eq(z1) [p(z2|z1)]

Marginalization:

q(z2 ∈ (a, b)) = Eq(z1,z2) [1(a < z2 < b)]

Action evaluation:Q(a) = Eq(s) [R(s, a)]

How to compute E [h(z)]?

if h(z) ≈∑i

αiγi(z)⇒ E [h(z)] ≈∑i

αiri

q(z)

(z)

, , , , , , , ,1 2 ... K[ ]r

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 10 / 20

Distributed distributional code

Why DDC? It makes computation simple for neurons

r := Eq(z) [γ1 (z) , γ2 (z) , ..., γK (z)]

Key computations involve expected values:

Message passing:q(z2) = Eq(z1) [p(z2|z1)]

Marginalization:

q(z2 ∈ (a, b)) = Eq(z1,z2) [1(a < z2 < b)]

Action evaluation:Q(a) = Eq(s) [R(s, a)]

How to compute E [h(z)]?

if h(z) ≈∑i

αiγi(z)⇒ E [h(z)] ≈∑i

αiri

q(z)h(z)

(z)

, , , , , , , ,1 2 ... K[ ]r

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 10 / 20

Distributed distributional code

Why DDC? It makes computation simple for neurons

r := Eq(z) [γ1 (z) , γ2 (z) , ..., γK (z)]

Key computations involve expected values:

Message passing:q(z2) = Eq(z1) [p(z2|z1)]

Marginalization:

q(z2 ∈ (a, b)) = Eq(z1,z2) [1(a < z2 < b)]

Action evaluation:Q(a) = Eq(s) [R(s, a)]

How to compute E [h(z)]?

if h(z) ≈∑i

αiγi(z)

⇒ E [h(z)] ≈∑i

αiri

q(z)h(z)

(z)

, , , , , , , ,1 2 ... K[ ]r

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 10 / 20

Distributed distributional code

Why DDC? It makes computation simple for neurons

r := Eq(z) [γ1 (z) , γ2 (z) , ..., γK (z)]

Key computations involve expected values:

Message passing:q(z2) = Eq(z1) [p(z2|z1)]

Marginalization:

q(z2 ∈ (a, b)) = Eq(z1,z2) [1(a < z2 < b)]

Action evaluation:Q(a) = Eq(s) [R(s, a)]

How to compute E [h(z)]?

if h(z) ≈∑i

αiγi(z)⇒ E [h(z)] ≈∑i

αiri

q(z)h(z)

(z)

, , , , , , , ,1 2 ... K[ ]r

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 10 / 20

Distributed distributional code

Why DDC? It allows biologically plausible learning to infer

Previously, we saw marginal q(z)↔ Eq(z) [γ(z)]

For p(z,x) = p(z)p(x|z), p(z|x) is intractable

How to obtain approx. q(z|x)↔ Eq(z|x) [γ(z)]?

Clue: posterior mean...

Ep(z|x) [γ(z)] = arg minφ

Ep(z|x)�‖γ(z)− φ‖22

�Amortize� using φW (x) := Wσ(x)

W ∗ = arg minW

Ep(z,x)�‖γ(z)−Wσ(x)‖22

r(x) := W ∗σ(x) = Eq(z|x) [γ(z)]

Find W ∗ by the delta rule:

∆W ∝ (γ(z)− φW (x))σ(x)ᵀ, {z,x} ∼ p

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 11 / 20

Distributed distributional code

Why DDC? It allows biologically plausible learning to infer

Previously, we saw marginal q(z)↔ Eq(z) [γ(z)]

For p(z,x) = p(z)p(x|z), p(z|x) is intractable

How to obtain approx. q(z|x)↔ Eq(z|x) [γ(z)]?

Clue: posterior mean...

Ep(z|x) [γ(z)] = arg minφ

Ep(z|x)�‖γ(z)− φ‖22

�Amortize� using φW (x) := Wσ(x)

W ∗ = arg minW

Ep(z,x)�‖γ(z)−Wσ(x)‖22

r(x) := W ∗σ(x) = Eq(z|x) [γ(z)]

Find W ∗ by the delta rule:

∆W ∝ (γ(z)− φW (x))σ(x)ᵀ, {z,x} ∼ p

z

x

γ(z)

simulation

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 11 / 20

Distributed distributional code

Why DDC? It allows biologically plausible learning to infer

Previously, we saw marginal q(z)↔ Eq(z) [γ(z)]

For p(z,x) = p(z)p(x|z), p(z|x) is intractable

How to obtain approx. q(z|x)↔ Eq(z|x) [γ(z)]?

Clue: posterior mean...

Ep(z|x) [γ(z)] = arg minφ

Ep(z|x)�‖γ(z)− φ‖22

�Amortize� using φW (x) := Wσ(x)

W ∗ = arg minW

Ep(z,x)�‖γ(z)−Wσ(x)‖22

r(x) := W ∗σ(x) = Eq(z|x) [γ(z)]

Find W ∗ by the delta rule:

∆W ∝ (γ(z)− φW (x))σ(x)ᵀ, {z,x} ∼ p

z

x

γ(z)

simulation

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 11 / 20

Distributed distributional code

Why DDC? It allows biologically plausible learning to infer

Previously, we saw marginal q(z)↔ Eq(z) [γ(z)]

For p(z,x) = p(z)p(x|z), p(z|x) is intractable

How to obtain approx. q(z|x)↔ Eq(z|x) [γ(z)]?

Clue: posterior mean...

Ep(z|x) [γ(z)] = arg minφ

Ep(z|x)�‖γ(z)− φ‖22

�Amortize� using φW (x) := Wσ(x)

W ∗ = arg minW

Ep(z,x)�‖γ(z)−Wσ(x)‖22

r(x) := W ∗σ(x) = Eq(z|x) [γ(z)]

Find W ∗ by the delta rule:

∆W ∝ (γ(z)− φW (x))σ(x)ᵀ, {z,x} ∼ p

z

x

γ(z)

simulation

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 11 / 20

Distributed distributional code

Why DDC? It allows biologically plausible learning to infer

Previously, we saw marginal q(z)↔ Eq(z) [γ(z)]

For p(z,x) = p(z)p(x|z), p(z|x) is intractable

How to obtain approx. q(z|x)↔ Eq(z|x) [γ(z)]?

Clue: posterior mean...

Ep(z|x) [γ(z)] = arg minφ

Ep(z|x)�‖γ(z)− φ‖22

�Amortize� using φW (x) := Wσ(x)

W ∗ = arg minW

Ep(z,x)�‖γ(z)−Wσ(x)‖22

r(x) := W ∗σ(x) = Eq(z|x) [γ(z)]

Find W ∗ by the delta rule:

∆W ∝ (γ(z)− φW (x))σ(x)ᵀ, {z,x} ∼ p

z

x

γ(z)

simulation

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 11 / 20

Distributed distributional code

Why DDC? It allows biologically plausible learning to infer

Previously, we saw marginal q(z)↔ Eq(z) [γ(z)]

For p(z,x) = p(z)p(x|z), p(z|x) is intractable

How to obtain approx. q(z|x)↔ Eq(z|x) [γ(z)]?

Clue: posterior mean...

Ep(z|x) [γ(z)] = arg minφ

Ep(z|x)�‖γ(z)− φ‖22

�Amortize� using φW (x) := Wσ(x)

W ∗ = arg minW

Ep(z,x)�‖γ(z)−Wσ(x)‖22

r(x) := W ∗σ(x) = Eq(z|x) [γ(z)]

Find W ∗ by the delta rule:

∆W ∝ (γ(z)− φW (x))σ(x)ᵀ, {z,x} ∼ p

z

x

γ(z)

simulation

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 11 / 20

Distributed distributional code

Why DDC? It allows biologically plausible learning to infer

Previously, we saw marginal q(z)↔ Eq(z) [γ(z)]

For p(z,x) = p(z)p(x|z), p(z|x) is intractable

How to obtain approx. q(z|x)↔ Eq(z|x) [γ(z)]?

Clue: posterior mean...

Ep(z|x) [γ(z)] = arg minφ

Ep(z|x)�‖γ(z)− φ‖22

�Amortize� using φW (x) := Wσ(x)

W ∗ = arg minW

Ep(z,x)�‖γ(z)−Wσ(x)‖22

r(x) := W ∗σ(x) = Eq(z|x) [γ(z)]

Find W ∗ by the delta rule:

∆W ∝ (γ(z)− φW (x))σ(x)ᵀ, {z,x} ∼ p

z

x

γ(z)

simulation

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 11 / 20

Distributed distributional code

Why DDC? It allows biologically plausible learning to infer

Previously, we saw marginal q(z)↔ Eq(z) [γ(z)]

For p(z,x) = p(z)p(x|z), p(z|x) is intractable

How to obtain approx. q(z|x)↔ Eq(z|x) [γ(z)]?

Clue: posterior mean...

Ep(z|x) [γ(z)] = arg minφ

Ep(z|x)�‖γ(z)− φ‖22

�Amortize� using φW (x) := Wσ(x)

W ∗ = arg minW

Ep(z,x)�‖γ(z)−Wσ(x)‖22

r(x) := W ∗σ(x) = Eq(z|x) [γ(z)]

Find W ∗ by the delta rule:

∆W ∝ (γ(z)− φW (x))σ(x)ᵀ, {z,x} ∼ p

z

x

γ(z)

simulation

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 11 / 20

Distributed distributional code

Why DDC? It allows biologically plausible learning to infer

Previously, we saw marginal q(z)↔ Eq(z) [γ(z)]

For p(z,x) = p(z)p(x|z), p(z|x) is intractable

How to obtain approx. q(z|x)↔ Eq(z|x) [γ(z)]?

Clue: posterior mean...

Ep(z|x) [γ(z)] = arg minφ

Ep(z|x)�‖γ(z)− φ‖22

�Amortize� using φW (x) := Wσ(x)

W ∗ = arg minW

Ep(z,x)�‖γ(z)−Wσ(x)‖22

r(x) := W ∗σ(x) = Eq(z|x) [γ(z)]

Find W ∗ by the delta rule:

∆W ∝ (γ(z)− φW (x))σ(x)ᵀ, {z,x} ∼ p

γ(z)

σ(x)

∆Wz

x

r

W

γ(z)

simulationlearning

to infer

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 11 / 20

Distributed distributional code

DDC Summary

De�nition

DDC of q(z) associated tuning functions γ(z) is r := Eq(z) [γ(z)]

MaxEnt interpretation

rE[γ(z)]

↼−−−−−−−−−−−−−−−−⇁γ(z),MaxEnt

q(z) ∝ exp [θ · γ(z)]

Expectation approximationh(z) ≈ α · γ(z) =⇒ E [h(z)] ≈ α · r

Learning to infer given p(z, x)

r(x) = Eq(z|x) [γ(z)] = W ∗σ(x), ∆W ∝ (γ − φW )σᵀ, {z,x} ∼ p(z,x)

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 12 / 20

Distributed distributional code

DDC Summary

De�nition

DDC of q(z) associated tuning functions γ(z) is r := Eq(z) [γ(z)]

MaxEnt interpretation

rE[γ(z)]

↼−−−−−−−−−−−−−−−−⇁γ(z),MaxEnt

q(z) ∝ exp [θ · γ(z)]

Expectation approximationh(z) ≈ α · γ(z) =⇒ E [h(z)] ≈ α · r

Learning to infer given p(z, x)

r(x) = Eq(z|x) [γ(z)] = W ∗σ(x), ∆W ∝ (γ − φW )σᵀ, {z,x} ∼ p(z,x)

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 12 / 20

Distributed distributional code

DDC Summary

De�nition

DDC of q(z) associated tuning functions γ(z) is r := Eq(z) [γ(z)]

MaxEnt interpretation

rE[γ(z)]

↼−−−−−−−−−−−−−−−−⇁γ(z),MaxEnt

q(z) ∝ exp [θ · γ(z)]

Expectation approximationh(z) ≈ α · γ(z) =⇒ E [h(z)] ≈ α · r

Learning to infer given p(z, x)

r(x) = Eq(z|x) [γ(z)] = W ∗σ(x), ∆W ∝ (γ − φW )σᵀ, {z,x} ∼ p(z,x)

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 12 / 20

Distributed distributional code

DDC Summary

De�nition

DDC of q(z) associated tuning functions γ(z) is r := Eq(z) [γ(z)]

MaxEnt interpretation

rE[γ(z)]

↼−−−−−−−−−−−−−−−−⇁γ(z),MaxEnt

q(z) ∝ exp [θ · γ(z)]

Expectation approximationh(z) ≈ α · γ(z) =⇒ E [h(z)] ≈ α · r

Learning to infer given p(z, x)

r(x) = Eq(z|x) [γ(z)] = W ∗σ(x), ∆W ∝ (γ − φW )σᵀ, {z,x} ∼ p(z,x)

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 12 / 20

Distributed distributional code

DDC Summary

De�nition

DDC of q(z) associated tuning functions γ(z) is r := Eq(z) [γ(z)]

MaxEnt interpretation

rE[γ(z)]

↼−−−−−−−−−−−−−−−−⇁γ(z),MaxEnt

q(z) ∝ exp [θ · γ(z)]

Expectation approximationh(z) ≈ α · γ(z) =⇒ E [h(z)] ≈ α · r

Learning to infer given p(z, x)

r(x) = Eq(z|x) [γ(z)] = W ∗σ(x), ∆W ∝ (γ − φW )σᵀ, {z,x} ∼ p(z,x)

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 12 / 20

Online recognition and postdiction

3. Online recognition and postdiction

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 13 / 20

Online recognition and postdiction

A generic dynamic internal model

We assume a generic internal model

zt = f(zt−1, ξ(z))

xt = g(zt, ξ(x))

Assumptions

Discrete-time

Markov property

Stationarity

zt−1 zt

xtxt−1

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 14 / 20

Online recognition and postdiction

A generic dynamic internal model

We assume a generic internal model

zt = f(zt−1, ξ(z))

xt = g(zt, ξ(x))

Assumptions

Discrete-time

Markov property

Stationarity

zt−1 zt

xtxt−1

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 14 / 20

Online recognition and postdiction

Representing and computing beliefs of the whole history

Online recognition (�ltering): maintain q(zt|x1:t)

De�ne temporally extended encoding function ψt := ψ (z1:t) for DDC

A plausible ψt

ψ1 := γ (z1)

ψt := Uψt−1 + γ (zt)

Update beliefs about the past: compute

rt = Eq(z1:t|x1:t) [ψt]

from rt−1 and xtPostiction: readout statistics

h(zt−τ ) ≈ α ·ψ(z1:t) =⇒ Eq(zt−τ ) [h(zt−τ )] ≈ α · rt

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 15 / 20

Online recognition and postdiction

Representing and computing beliefs of the whole history

Online recognition (�ltering): maintain q(z1:t|x1:t) to allow postdiction

De�ne temporally extended encoding function ψt := ψ (z1:t) for DDC

A plausible ψt

ψ1 := γ (z1)

ψt := Uψt−1 + γ (zt)

Update beliefs about the past: compute

rt = Eq(z1:t|x1:t) [ψt]

from rt−1 and xtPostiction: readout statistics

h(zt−τ ) ≈ α ·ψ(z1:t) =⇒ Eq(zt−τ ) [h(zt−τ )] ≈ α · rt

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 15 / 20

Online recognition and postdiction

Representing and computing beliefs of the whole history

Online recognition (�ltering): maintain q(z1:t|x1:t) to allow postdiction

De�ne temporally extended encoding function ψt := ψ (z1:t) for DDC

A plausible ψt

ψ1 := γ (z1)

ψt := Uψt−1 + γ (zt)

Update beliefs about the past: compute

rt = Eq(z1:t|x1:t) [ψt]

from rt−1 and xtPostiction: readout statistics

h(zt−τ ) ≈ α ·ψ(z1:t) =⇒ Eq(zt−τ ) [h(zt−τ )] ≈ α · rt

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 15 / 20

Online recognition and postdiction

Representing and computing beliefs of the whole history

Online recognition (�ltering): maintain q(z1:t|x1:t) to allow postdiction

De�ne temporally extended encoding function ψt := ψ (z1:t) for DDC

A plausible ψt

ψ1 := γ (z1)

ψt := Uψt−1 + γ (zt)

Update beliefs about the past: compute

rt = Eq(z1:t|x1:t) [ψt]

from rt−1 and xtPostiction: readout statistics

h(zt−τ ) ≈ α ·ψ(z1:t) =⇒ Eq(zt−τ ) [h(zt−τ )] ≈ α · rt

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 15 / 20

Online recognition and postdiction

Representing and computing beliefs of the whole history

Online recognition (�ltering): maintain q(z1:t|x1:t) to allow postdiction

De�ne temporally extended encoding function ψt := ψ (z1:t) for DDC

A plausible ψt

ψ1 := γ (z1)

ψt := Uψt−1 + γ (zt)

Update beliefs about the past: compute

rt = Eq(z1:t|x1:t) [ψt]

from rt−1 and xt

Postiction: readout statistics

h(zt−τ ) ≈ α ·ψ(z1:t) =⇒ Eq(zt−τ ) [h(zt−τ )] ≈ α · rt

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 15 / 20

Online recognition and postdiction

Representing and computing beliefs of the whole history

Online recognition (�ltering): maintain q(z1:t|x1:t) to allow postdiction

De�ne temporally extended encoding function ψt := ψ (z1:t) for DDC

A plausible ψt

ψ1 := γ (z1)

ψt := Uψt−1 + γ (zt)

Update beliefs about the past: compute

rt = Eq(z1:t|x1:t) [ψt]

from rt−1 and xtPostiction: readout statistics

h(zt−τ ) ≈ α ·ψ(z1:t) =⇒ Eq(zt−τ ) [h(zt−τ )] ≈ α · rt

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 15 / 20

Online recognition and postdiction

Learning to postdict online

Want rt(x1:t) = Eqt [ψt (z1:t)]

Recall for p(z,x)r := φW ∗(x)

Likewise, for SSM p(z1:t,x1:t)

rt := φW t(x1:t)

Learning by the delta rule

∆W t ← (ψt − φt)(rt−1 ⊗ σt)ᵀ

{ψt,xt, rt−1} ∼ p(z1:t,x1:t), {hW i}t−1i=1

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 16 / 20

Online recognition and postdiction

Learning to postdict online

Want rt(x1:t) = Eqt [ψt (z1:t)]

Recall for p(z,x)r := φW ∗(x)

Likewise, for SSM p(z1:t,x1:t)

rt := φW t(x1:t)

Learning by the delta rule

∆W t ← (ψt − φt)(rt−1 ⊗ σt)ᵀ

{ψt,xt, rt−1} ∼ p(z1:t,x1:t), {hW i}t−1i=1

γ(z)

σ(x)

∆Wz

x

r

W

γ(z)

simulationlearning

to infer

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 16 / 20

Online recognition and postdiction

Learning to postdict online

Want rt(x1:t) = Eqt [ψt (z1:t)]

Recall for p(z,x)r := φW ∗(x)

Likewise, for SSM p(z1:t,x1:t)

rt := φW t(x1:t)

Learning by the delta rule

∆W t ← (ψt − φt)(rt−1 ⊗ σt)ᵀ

{ψt,xt, rt−1} ∼ p(z1:t,x1:t), {hW i}t−1i=1

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 16 / 20

Online recognition and postdiction

Learning to postdict online

Want rt(x1:t) = Eqt [ψt (z1:t)]

Recall for p(z,x)r := φW ∗(x)

Likewise, for SSM p(z1:t,x1:t)

rt := φW t(rt−1,xt)

Learning by the delta rule

∆W t ← (ψt − φt)(rt−1 ⊗ σt)ᵀ

{ψt,xt, rt−1} ∼ p(z1:t,x1:t), {hW i}t−1i=1

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 16 / 20

Online recognition and postdiction

Learning to postdict online

Want rt(x1:t) = Eqt [ψt (z1:t)]

Recall for p(z,x)r := φW ∗(x)

Likewise, for SSM p(z1:t,x1:t)

rt := φW t(rt−1,xt)

= W t · (rt−1 ⊗ σ(xt))

Learning by the delta rule

∆W t ← (ψt − φt)(rt−1 ⊗ σt)ᵀ

{ψt,xt, rt−1} ∼ p(z1:t,x1:t), {hW i}t−1i=1

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 16 / 20

Online recognition and postdiction

Learning to postdict online

Want rt(x1:t) = Eqt [ψt (z1:t)]

Recall for p(z,x)r := φW ∗(x)

Likewise, for SSM p(z1:t,x1:t)

rt := φW t(rt−1,xt)

= W t · (rt−1 ⊗ σ(xt))

Learning by the delta rule

∆W t ← (ψt − φt)(rt−1 ⊗ σt)ᵀ

{ψt,xt, rt−1} ∼ p(z1:t,x1:t), {hW i}t−1i=1

ψt

σ(xt)

rt−1

∆W tzt−1 zt

xt

rt

W t

rt−1

W t+1

ψtψt−1

Simulation Learning to infer

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 16 / 20

Online recognition and postdiction

Learning to postdict online

Want rt(x1:t) = Eqt [ψt (z1:t)]

Recall for p(z,x)r := φW ∗(x)

Likewise, for SSM p(z1:t,x1:t)

rt := φW t(rt−1,xt)

= W t · (rt−1 ⊗ σ(xt))

Learning by the delta rule

∆W t ← (ψt − φt)(rt−1 ⊗ σt)ᵀ

{ψt,xt, rt−1} ∼ p(z1:t,x1:t), {hW i}t−1i=1

ψt

σ(xt)

rt−1

∆W tzt−1 zt

xt

rt

W t

rt−1

W t+1

ψtψt−1

Simulation Learning to infer

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 16 / 20

Online recognition and postdiction

Learning to postdict online

Want rt(x1:t) = Eqt [ψt (z1:t)]

Recall for p(z,x)r := φW ∗(x)

Likewise, for SSM p(z1:t,x1:t)

rt := φW t(rt−1,xt)

= W t · (rt−1 ⊗ σ(xt))

Learning by the delta rule

∆W t ← (ψt − φt)(rt−1 ⊗ σt)ᵀ

{ψt,xt, rt−1} ∼ p(z1:t,x1:t), {hW i}t−1i=1

ψt

σ(xt)

rt−1

∆W tzt−1 zt

xt

rt

W t

rt−1

W t+1

ψtψt−1

Simulation Learning to infer

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 16 / 20

Testing DDC �ltering on simulated experiments

4. Testing DDC �ltering on simulated experiments

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 17 / 20

Testing DDC �ltering on simulated experiments

Auditory continuity illusion

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 18 / 20

Testing DDC �ltering on simulated experiments

Auditory continuity illusion

0

5st

ate

true states of tone and noisetone noise

0 50time

freq

spectrogram observations

0 4

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 18 / 20

Testing DDC �ltering on simulated experiments

Auditory continuity illusion

stat

e tone mask

freq

1 2 3 4 5 6 7 8 9 10stimulus time (t− τ)

10987654321

perc

eptio

n tim

e (t)

0 1

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 18 / 20

Testing DDC �ltering on simulated experiments

Auditory continuity illusion

stat

e tone mask

freq

1 2 3 4 5 6 7 8 9 10stimulus time (t− τ)

10987654321

perc

eptio

n tim

e (t)

0 1

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 18 / 20

Testing DDC �ltering on simulated experiments

Auditory continuity illusion

stat

e tone mask

freq

1 2 3 4 5 6 7 8 9 10stimulus time (t− τ)

10987654321

perc

eptio

n tim

e (t)

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 18 / 20

Testing DDC �ltering on simulated experiments

Auditory continuity illusion

stat

e tone mask

freq

1 2 3 4 5 6 7 8 9 10stimulus time (t− τ)

10987654321

perc

eptio

n tim

e (t)

0 1

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 18 / 20

Testing DDC �ltering on simulated experiments

Auditory continuity illusion

stat

e tone mask

freq

1 2 3 4 5 6 7 8 9 10stimulus time (t− τ)

10987654321

perc

eptio

n tim

e (t)

0 1

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 18 / 20

Testing DDC �ltering on simulated experiments

Auditory continuity illusion

stat

e tone mask

freq

1 2 3 4 5 6 7 8 9 10stimulus time (t− τ)

10987654321

perc

eptio

n tim

e (t)

0 1

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 18 / 20

Testing DDC �ltering on simulated experiments

Auditory continuity illusion

stat

e tone mask

freq

1 2 3 4 5 6 7 8 9 10stimulus time (t− τ)

10987654321

perc

eptio

n tim

e (t)

0 1

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 18 / 20

Testing DDC �ltering on simulated experiments

Auditory continuity illusion

stat

e tone mask

freq

1 2 3 4 5 6 7 8 9 10stimulus time (t− τ)

10987654321

perc

eptio

n tim

e (t)

0 1

stat

e tone mask

freq

1 2 3 4 5 6 7 8 9 10stimulus time (t− τ)

10987654321

perc

eptio

n tim

e (t)

0 1

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 18 / 20

Testing DDC �ltering on simulated experiments

Auditory continuity illusion

stat

e tone mask

freq

1 2 3 4 5 6 7 8 9 10stimulus time (t− τ)

10987654321

perc

eptio

n tim

e (t)

0 1

stat

e tone mask

freq

1 2 3 4 5 6 7 8 9 10stimulus time (t− τ)

10987654321

perc

eptio

n tim

e (t)

0 1

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 18 / 20

Testing DDC �ltering on simulated experiments

Auditory continuity illusion

stat

e tone mask

freq

1 2 3 4 5 6 7 8 9 10stimulus time (t− τ)

10987654321

perc

eptio

n tim

e (t)

0 1

stat

e tone mask

freq

1 2 3 4 5 6 7 8 9 10stimulus time (t− τ)

10987654321

perc

eptio

n tim

e (t)

0 1

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 18 / 20

Testing DDC �ltering on simulated experiments

Auditory continuity illusion

stat

e tone mask

freq

1 2 3 4 5 6 7 8 9 10stimulus time (t− τ)

10987654321

perc

eptio

n tim

e (t)

0 1

stat

e tone mask

freq

1 2 3 4 5 6 7 8 9 10stimulus time (t− τ)

10987654321

perc

eptio

n tim

e (t)

0 1

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 18 / 20

Testing DDC �ltering on simulated experiments

Auditory continuity illusion

stat

e tone mask

freq

1 2 3 4 5 6 7 8 9 10stimulus time (t− τ)

10987654321

perc

eptio

n tim

e (t)

0 1

stat

e tone mask

freq

1 2 3 4 5 6 7 8 9 10stimulus time (t− τ)

10987654321

perc

eptio

n tim

e (t)

0 1

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 18 / 20

Testing DDC �ltering on simulated experiments

Auditory continuity illusion

stat

e tone mask

freq

1 2 3 4 5 6 7 8 9 10stimulus time (t− τ)

10987654321

perc

eptio

n tim

e (t)

0 1

stat

e tone mask

freq

1 2 3 4 5 6 7 8 9 10stimulus time (t− τ)

10987654321

perc

eptio

n tim

e (t)

0 1

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 18 / 20

Testing DDC �ltering on simulated experiments

Auditory continuity illusion

stat

e tone mask

freq

1 2 3 4 5 6 7 8 9 10stimulus time (t− τ)

10987654321

perc

eptio

n tim

e (t)

0 1

stat

e tone mask

freq

1 2 3 4 5 6 7 8 9 10stimulus time (t− τ)

10987654321

perc

eptio

n tim

e (t)

0 1

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 18 / 20

Testing DDC �ltering on simulated experiments

Auditory continuity illusion

stat

e tone mask

freq

1 2 3 4 5 6 7 8 9 10stimulus time (t− τ)

10987654321

perc

eptio

n tim

e (t)

0 1

stat

e tone mask

freq

1 2 3 4 5 6 7 8 9 10stimulus time (t− τ)

10987654321

perc

eptio

n tim

e (t)

0 1

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 18 / 20

Testing DDC �ltering on simulated experiments

Auditory continuity illusion

stat

e tone mask

freq

1 2 3 4 5 6 7 8 9 10stimulus time (t− τ)

10987654321

perc

eptio

n tim

e (t)

0 1

stat

e tone mask

freq

1 2 3 4 5 6 7 8 9 10stimulus time (t− τ)

10987654321

perc

eptio

n tim

e (t)

0 1

stat

e tone mask

freq

1 2 3 4 5 6 7 8 9 10stimulus time (t− τ)

10987654321

perc

eptio

n tim

e (t)

0 1

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 18 / 20

Testing DDC �ltering on simulated experiments

Auditory continuity illusion

stat

e tone mask

freq

1 2 3 4 5 6 7 8 9 10stimulus time (t− τ)

10987654321

perc

eptio

n tim

e (t)

0 1

stat

e tone mask

freq

1 2 3 4 5 6 7 8 9 10stimulus time (t− τ)

10987654321

perc

eptio

n tim

e (t)

0 1

stat

e tone mask

freq

1 2 3 4 5 6 7 8 9 10stimulus time (t− τ)

10987654321

perc

eptio

n tim

e (t)

0 1

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 18 / 20

Testing DDC �ltering on simulated experiments

Auditory continuity illusion

stat

e tone mask

freq

1 2 3 4 5 6 7 8 9 10stimulus time (t− τ)

10987654321

perc

eptio

n tim

e (t)

0 1

stat

e tone mask

freq

1 2 3 4 5 6 7 8 9 10stimulus time (t− τ)

10987654321

perc

eptio

n tim

e (t)

0 1

stat

e tone mask

freq

1 2 3 4 5 6 7 8 9 10stimulus time (t− τ)

10987654321

perc

eptio

n tim

e (t)

0 1

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 18 / 20

Testing DDC �ltering on simulated experiments

Auditory continuity illusion

stat

e tone mask

freq

1 2 3 4 5 6 7 8 9 10stimulus time (t− τ)

10987654321

perc

eptio

n tim

e (t)

0 1

stat

e tone mask

freq

1 2 3 4 5 6 7 8 9 10stimulus time (t− τ)

10987654321

perc

eptio

n tim

e (t)

0 1

stat

e tone mask

freq

1 2 3 4 5 6 7 8 9 10stimulus time (t− τ)

10987654321

perc

eptio

n tim

e (t)

0 1

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 18 / 20

Testing DDC �ltering on simulated experiments

Cutaneous rabbit

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 19 / 20

Testing DDC �ltering on simulated experiments

Cutaneous rabbit

1

0

1

loca

tion

true tap locations

0 20 40 60 80# taps

30

1

neur

on

spiking observations

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 19 / 20

Testing DDC �ltering on simulated experiments

Cutaneous rabbit

1 6 11 15# taps

0

1

2

posit

ion

true tap

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 19 / 20

Testing DDC �ltering on simulated experiments

Cutaneous rabbit

1 6 11 15# taps

0

1

2

trueperceived

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 19 / 20

Testing DDC �ltering on simulated experiments

Cutaneous rabbit

1 6 11 15# taps

0

1

2

trueperceived

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 19 / 20

Testing DDC �ltering on simulated experiments

Cutaneous rabbit

1 6 11 15# taps

0

1

2

trueperceived

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 19 / 20

Testing DDC �ltering on simulated experiments

Cutaneous rabbit

1 6 11 15# taps

0

1

2

trueperceived

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 19 / 20

Testing DDC �ltering on simulated experiments

Cutaneous rabbit

1 6 11 15# taps

0

1

2

trueperceived

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 19 / 20

Testing DDC �ltering on simulated experiments

Cutaneous rabbit

1 6 11 15# taps

0

1

2

trueperceived

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 19 / 20

Testing DDC �ltering on simulated experiments

Cutaneous rabbit

1 6 11 15# taps

0

1

2

trueperceived

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 19 / 20

Testing DDC �ltering on simulated experiments

Cutaneous rabbit

1 6 11 15# taps

0

1

2

trueperceived

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 19 / 20

Testing DDC �ltering on simulated experiments

Cutaneous rabbit

1 6 11 15# taps

0

1

2

trueperceived

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 19 / 20

Testing DDC �ltering on simulated experiments

Cutaneous rabbit

1 6 11 15# taps

0

1

2

trueperceived

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 19 / 20

Testing DDC �ltering on simulated experiments

Cutaneous rabbit

1 6 11 15# taps

0

1

2

trueperceived

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 19 / 20

Testing DDC �ltering on simulated experiments

Cutaneous rabbit

1 6 11 15# taps

0

1

2

trueperceived

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 19 / 20

Testing DDC �ltering on simulated experiments

Cutaneous rabbit

1 6 11 15# taps

0

1

2

trueperceived

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 19 / 20

Testing DDC �ltering on simulated experiments

Cutaneous rabbit

1 6 11 15# taps

0

1

2

trueperceived

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 19 / 20

Testing DDC �ltering on simulated experiments

Cutaneous rabbit

1 6 11 15# taps

0

1

2

trueperceived

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 19 / 20

Testing DDC �ltering on simulated experiments

Cutaneous rabbit

1 6 11 15# taps

0

1

2

trueperceived

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 19 / 20

Testing DDC �ltering on simulated experiments

Summary of DDC �ltering and Questions

Representation: DDC rt := Eq(z1:t|x1:t) [ψt(z1:t)]

Computation: bilinear rt := W ∗ · (rt−1 ⊗ σ(xt))

, or linear Eq [h(zt−τ )] ≈ αᵀrt

Learning to infer: delta rule ∆W ∝ (ψt − φW )(rt−1 ⊗ σt), similar for readout αQuestions:

Is the encoding function ψt = Uψt−1 + γ(zt) the best?

Does the brain encode the joint q(z1:t|x1:t)?

Any theoretical argument for using the bilinear rule?

Is there a way to also adapt U in a plausible way?

Can we encode the internal model by DDC? (talk to Eszter Vértes)

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 20 / 20

Testing DDC �ltering on simulated experiments

Summary of DDC �ltering and Questions

Representation: DDC rt := Eq(z1:t|x1:t) [ψt(z1:t)]Computation: bilinear rt := W ∗ · (rt−1 ⊗ σ(xt)) , or linear Eq [h(zt−τ )] ≈ αᵀrt

Learning to infer: delta rule ∆W ∝ (ψt − φW )(rt−1 ⊗ σt), similar for readout αQuestions:

Is the encoding function ψt = Uψt−1 + γ(zt) the best?

Does the brain encode the joint q(z1:t|x1:t)?

Any theoretical argument for using the bilinear rule?

Is there a way to also adapt U in a plausible way?

Can we encode the internal model by DDC? (talk to Eszter Vértes)

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 20 / 20

Testing DDC �ltering on simulated experiments

Summary of DDC �ltering and Questions

Representation: DDC rt := Eq(z1:t|x1:t) [ψt(z1:t)]Computation: bilinear rt := W ∗ · (rt−1 ⊗ σ(xt)) , or linear Eq [h(zt−τ )] ≈ αᵀrtLearning to infer: delta rule ∆W ∝ (ψt − φW )(rt−1 ⊗ σt), similar for readout α

Questions:

Is the encoding function ψt = Uψt−1 + γ(zt) the best?

Does the brain encode the joint q(z1:t|x1:t)?

Any theoretical argument for using the bilinear rule?

Is there a way to also adapt U in a plausible way?

Can we encode the internal model by DDC? (talk to Eszter Vértes)

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 20 / 20

Testing DDC �ltering on simulated experiments

Summary of DDC �ltering and Questions

Representation: DDC rt := Eq(z1:t|x1:t) [ψt(z1:t)]Computation: bilinear rt := W ∗ · (rt−1 ⊗ σ(xt)) , or linear Eq [h(zt−τ )] ≈ αᵀrtLearning to infer: delta rule ∆W ∝ (ψt − φW )(rt−1 ⊗ σt), similar for readout αQuestions:

Is the encoding function ψt = Uψt−1 + γ(zt) the best?

Does the brain encode the joint q(z1:t|x1:t)?

Any theoretical argument for using the bilinear rule?

Is there a way to also adapt U in a plausible way?

Can we encode the internal model by DDC? (talk to Eszter Vértes)

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 20 / 20

Testing DDC �ltering on simulated experiments

Summary of DDC �ltering and Questions

Representation: DDC rt := Eq(z1:t|x1:t) [ψt(z1:t)]Computation: bilinear rt := W ∗ · (rt−1 ⊗ σ(xt)) , or linear Eq [h(zt−τ )] ≈ αᵀrtLearning to infer: delta rule ∆W ∝ (ψt − φW )(rt−1 ⊗ σt), similar for readout αQuestions:

Is the encoding function ψt = Uψt−1 + γ(zt) the best?

Does the brain encode the joint q(z1:t|x1:t)?

Any theoretical argument for using the bilinear rule?

Is there a way to also adapt U in a plausible way?

Can we encode the internal model by DDC? (talk to Eszter Vértes)

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 20 / 20

Testing DDC �ltering on simulated experiments

Summary of DDC �ltering and Questions

Representation: DDC rt := Eq(z1:t|x1:t) [ψt(z1:t)]Computation: bilinear rt := W ∗ · (rt−1 ⊗ σ(xt)) , or linear Eq [h(zt−τ )] ≈ αᵀrtLearning to infer: delta rule ∆W ∝ (ψt − φW )(rt−1 ⊗ σt), similar for readout αQuestions:

Is the encoding function ψt = Uψt−1 + γ(zt) the best?

Does the brain encode the joint q(z1:t|x1:t)?

Any theoretical argument for using the bilinear rule?

Is there a way to also adapt U in a plausible way?

Can we encode the internal model by DDC? (talk to Eszter Vértes)

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 20 / 20

Testing DDC �ltering on simulated experiments

Summary of DDC �ltering and Questions

Representation: DDC rt := Eq(z1:t|x1:t) [ψt(z1:t)]Computation: bilinear rt := W ∗ · (rt−1 ⊗ σ(xt)) , or linear Eq [h(zt−τ )] ≈ αᵀrtLearning to infer: delta rule ∆W ∝ (ψt − φW )(rt−1 ⊗ σt), similar for readout αQuestions:

Is the encoding function ψt = Uψt−1 + γ(zt) the best?

Does the brain encode the joint q(z1:t|x1:t)?

Any theoretical argument for using the bilinear rule?

Is there a way to also adapt U in a plausible way?

Can we encode the internal model by DDC? (talk to Eszter Vértes)

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 20 / 20

Testing DDC �ltering on simulated experiments

Summary of DDC �ltering and Questions

Representation: DDC rt := Eq(z1:t|x1:t) [ψt(z1:t)]Computation: bilinear rt := W ∗ · (rt−1 ⊗ σ(xt)) , or linear Eq [h(zt−τ )] ≈ αᵀrtLearning to infer: delta rule ∆W ∝ (ψt − φW )(rt−1 ⊗ σt), similar for readout αQuestions:

Is the encoding function ψt = Uψt−1 + γ(zt) the best?

Does the brain encode the joint q(z1:t|x1:t)?

Any theoretical argument for using the bilinear rule?

Is there a way to also adapt U in a plausible way?

Can we encode the internal model by DDC? (talk to Eszter Vértes)

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 20 / 20

Testing DDC �ltering on simulated experiments

Summary of DDC �ltering and Questions

Representation: DDC rt := Eq(z1:t|x1:t) [ψt(z1:t)]Computation: bilinear rt := W ∗ · (rt−1 ⊗ σ(xt)) , or linear Eq [h(zt−τ )] ≈ αᵀrtLearning to infer: delta rule ∆W ∝ (ψt − φW )(rt−1 ⊗ σt), similar for readout αQuestions:

Is the encoding function ψt = Uψt−1 + γ(zt) the best?

Does the brain encode the joint q(z1:t|x1:t)?

Any theoretical argument for using the bilinear rule?

Is there a way to also adapt U in a plausible way?

Can we encode the internal model by DDC? (talk to Eszter Vértes)

Kevin Li, Maneesh Sahani (Gatsby Unit, UCL) Model of recognition and postdiction January 6, 2020 20 / 20

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