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How else can we define Information Flow in Neural Circuits? Praveen Venkatesh with Sanghamitra Dutta and Pulkit Grover Dept. of Electrical & Computer Engineering Carnegie Mellon University How should we define Information Flow in Neural


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How else can we define Information Flow in Neural Circuits?

“How should we define Information Flow in Neural Circuits?”, ISIT 2019 “Information Flow in Computational Systems”, IEEE Trans. IT 2020 (https://praveenv253.github.io/publications)

with Sanghamitra Dutta and Pulkit Grover

Praveen Venkatesh

  • Dept. of Electrical & Computer Engineering

Carnegie Mellon University

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Acknowledgments

Theorists Neuroscientists Clinicians Labmates & Colleagues Advisor

Pulkit Grover (CMU) Marlene Behrmann (CMU) Rob Kass (CMU) Mark Richardson (MGH/Harvard) Vasily Kokkinos (MGH/Hvd) Bobak Nazer (BU) Venkatesh Saligrama (BU) Sanghamitra Dutta (CMU) Aditya Gangrade (BU) Haewon Jeong (CMU) Ashwati Krishnan (CMU) Alireza Chamanzar (CMU)

Fellowships

  • CMLH Fellowship in Digital Health
  • Dowd Fellowship
  • Henry L. Hillman Presidential Fellowship
  • CIT Dean’s Fellowship

Todd Coleman (UCSD)

Grants

  • Center for Machine Learning and Health
  • Chuck Noll Foundation for Brain Injury

Research

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  • Thousands of papers estimating information flow in the brain
  • Lot of controversy around information flow

Our main insight: controversy exists because “information flow” is not formally defined

(Venkatesh & Grover SfN’15, Allerton’15) (Venkatesh & Grover, Cosyne’19) (Venkatesh, Dutta & Grover, IEEE Trans IT ’20)

Why study information flow?

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4

How else can we define Information Flow?

  • 1. Why study information flow?
  • 2. ISIT’19 recap: How should we define Information Flow?
  • Shortcoming: “𝑁-information Orphans”
  • Potential Fix based on Pruning
  • 3. Information Flow from a Causality Perspective
  • A counterexample to Pruning
  • Intro to Causality and Counterfactual Causal Influence
  • 4. CCI is the intuitive definition, but not observational
  • An Alternative Observational Definition
  • A Comparison of Definitions
  • 5. Conclusion
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5

(Almeida et al., Cortex, 2013)

Region A Region C Region B “Stimulus” Goal: Find a definition for information flow, so that we can track info paths

  • Info “flows” between

brain regions

  • Info is often about a

stimulus

  • There could be feedback

Information flow for Neuroscientific Inferences

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6 𝑁: “Message” = stimulus 𝑌(𝐹𝑢): Transmission on edge 𝐹𝑢 at time 𝑢

  • “Brain areas”
  • Feedback communication
  • 𝑌(𝐹0) is the transmission on

the edge 𝐹0 at 𝑢 = 0 𝑌(𝐹1) = 𝑔

𝐵1 𝑌 𝐹0 , …

  • Transmissions on edges are

measured

  • Message (a.k.a. stimulus)

arrives at and only at 𝑢 = 0

(Thompson, 1980: VLSI) (Ahlswede et al., 2000: Network Info Theory) (Peters et al., 2016: Causality)

𝐵0 𝐶0 𝐷0

𝑢 = 0

𝐵1 𝐶1 𝐷1

𝑢 = 1

𝐵2 𝐶2 𝐷2

𝑢 = 2 𝑁 𝑔(𝑁) 𝑌(𝐹1) 𝑌(𝐹0)

Region A Region C Region B Stimulus

A Computational Model of the Brain

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7

  • “Brain areas”
  • Feedback communication
  • 𝑌(𝐹0) is the transmission on

the edge 𝐹0 at 𝑢 = 0 𝑌(𝐹1) = 𝑔

𝐵1 𝑌 𝐹0 , …

  • Transmissions on edges are

measured

  • Message (a.k.a. stimulus)

arrives at and only at 𝑢 = 0

(Thompson, 1980: VLSI) (Ahlswede et al., 2000: Network Info Theory) (Peters et al., 2016: Causality)

𝐵0 𝐵1 𝐵2 𝐶0 𝐶1 𝐶2

𝑁 𝑔(𝑁)

𝐷0 𝐷1 𝐷2

𝑌(𝐹0) 𝑌(𝐹1) 𝑢 = 0 𝑢 = 1 𝑢 = 2

Goal: Define info flow + track info path

A Computational Model of the Brain

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𝑁, 𝑎 ~ iid Ber(1/2)

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𝑁 ⊕ 𝑎 = 𝑁 xor 𝑎 𝐽 𝑁; 𝑁 ⊕ 𝑎 = 0 Why not a simpler definition like: 𝐽 𝑁; 𝑌 𝐹𝑢 > 0

?

Definition [𝑁-Information Flow]: (ISIT ’19, Trans IT ‘20) We say that an edge 𝐹𝑢 has 𝑁-information flow if ∃ ℰ𝑢

′ ⊆ ℰ𝑢 s.t. 𝐽 𝑁; 𝑌 𝐹𝑢 | 𝑌(ℰ𝑢 ′) > 0.

How should we define Information Flow? (ISIT ‘19)

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Using the above definition: 𝐽 𝑁; 𝑁 ⊕ 𝑎 𝑎 > 0 Conditioning on 𝑎 reveals the dependence between 𝑁 ⊕ 𝑎 and 𝑁: Definition [𝑁-Information Flow]: (ISIT ‘19, Trans IT ‘20) We say that an edge 𝐹𝑢 has 𝑁-information flow if ∃ ℰ𝑢

′ ⊆ ℰ𝑢 s.t. 𝐽 𝑁; 𝑌 𝐹𝑢 | 𝑌(ℰ𝑢 ′) > 0.

How should we define Information Flow? (ISIT ‘19)

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Definition [𝑁-Information Flow]: (ISIT ’19, Trans IT ‘20) We say that an edge 𝐹𝑢 has 𝑁-information flow if ∃ ℰ𝑢

′ ⊆ ℰ𝑢 s.t. 𝐽 𝑁; 𝑌 𝐹𝑢 | 𝑌(ℰ𝑢 ′) > 0.

How should we define Information Flow? (ISIT ‘19)

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Theorem [Existence of Information Paths]: If the transmissions of an “output” node 𝑊

𝑢 𝑝𝑞 depend on 𝑁, then there

exists a path from 𝑊

𝑗𝑞 to 𝑊 𝑢 𝑝𝑞, every edge of which has 𝑁-info flow. (Venkatesh, Dutta & Grover, ISIT 2019; Trans. IEEE 2020)

𝐵0 𝐵1 𝐵2 𝐶0 𝐶1 𝐶2

𝑁 𝑔 𝑁 𝑌(𝐹0) 𝑌(𝐹1) 𝑊

𝑗𝑞

𝑊

𝑢 𝑝𝑞

The Existence of Information Paths

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An edge 𝐹𝑢 has 𝑁-info flow if ∃ ℰ𝑢

′ ⊆ ℰ𝑢 s.t. 𝐽 𝑁; 𝑌 𝐹𝑢 | 𝑌(ℰ𝑢 ′) > 0.

𝑁-info flows out of a node ⇒ 𝑁-info flows into the node 𝐽 𝑁; 𝑁 ⊕ 𝑎 𝑎 > 0 𝐽 𝑁; 𝑎 𝑁 ⊕ 𝑎 > 0 𝐷1 is an orphan 𝑎 has 𝑁-info flow!

But! This definition gives rise to “Orphans”

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Algorithm: Prune edges that do not lead to 𝑊

𝑗𝑞 using Depth First Search (Venkatesh, Dutta & Grover, IEEE Trans. IT 2020; to appear)

Pruning as a Solution to Orphans

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Algorithm: Prune edges that do not lead to 𝑊

𝑗𝑞 using Depth First Search (Venkatesh, Dutta & Grover, IEEE Trans. IT 2020; to appear)

Pruning as a Solution to Orphans

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Algorithm: Prune edges that do not lead to 𝑊

𝑗𝑞 using Depth First Search (Venkatesh, Dutta & Grover, IEEE Trans. IT 2020; to appear)

Pruning as a Solution to Orphans

Remove orphans such as 𝐷1 Hopefully prune out edges like 𝑎, which are not “computed” from 𝑁

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How else can we define Information Flow?

  • 1. Why study information flow?
  • 2. ISIT’19 recap: Defining information flow
  • Shortcoming: “𝑁-information Orphans”
  • Potential Fix based on Pruning
  • 3. Information Flow from a Causality Perspective
  • A counterexample to Pruning
  • Intro to Causality and Counterfactual Causal Influence
  • 4. CCI is the intuitive definition, but not observational
  • An Alternative Observational Definition
  • A Comparison of Definitions
  • 5. Conclusion
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𝐽 𝑁; 𝑁 ⊕ 𝑎 = 0 𝐽 𝑁; 𝑁 ⊕ 𝑎 | [𝑁, 𝑎] = 0 The only information path from 𝐵0 to 𝐶3 is through an edge carrying 𝑎! The orphan removed by pruning transmits 𝑁 ⊕ 𝑎! (𝐵1, 𝐵2) has no 𝑁-info flow:

𝑁, 𝑎 ~ Ber(1/2) 𝑁 ⊥ 𝑎

𝐵2 is an 𝑁-information Orphan

A Counterexample to Pruning (this work)

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We disliked orphans for two reasons: 1. No “conservation” of info flow at an orphan 2. 𝑎 is not “computed” from 𝑁 Is 𝑎 really all that different from 𝑁 ⊕ 𝑎 in these examples? To differentiate, we need to go beyond the joint distribution! 𝑞 𝑛, 𝑨, 𝑧 = 1 4 𝕁 𝑧 = 𝑛 ⊕ 𝑨 = 1 4 𝕁 𝑧 ⊕ 𝑨 = 𝑛 Using 𝑍 for 𝑁 ⊕ 𝑎, Joint distribution is symmetric in 𝑁 ⊕ 𝑎 and 𝑎!

What makes this a Counterexample?

𝑁, 𝑎 ~ Ber(1/2) 𝑁 ⊥ 𝑎

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19 𝑌0 𝑌1 𝑌4 𝑌2 𝑌5 𝑌3 Directed Acyclic Graph with functional relationships

𝑌𝑗 = 𝑔

𝑗 𝑄𝑏𝑌𝑗, 𝑋 𝑗

𝐵0 𝐶0 𝐵1 𝐶1 𝐵2 𝐶2

𝑁 𝑔(𝑁) 𝑌1 𝑌2 𝑌3 𝑌3 = 𝑔

𝐵1 𝑌1, 𝑌2, 𝑋 𝐵1

The Computational System is also a Structural Causal Model Structural Causal Model

(Peters, Janzing & Schölkopf, “Elements of Causal Inference”, 2016)

Computational System Model

(Venkatesh, Dutta & Grover, ISIT 2019; Trans. IT 2020)

A Brief Introduction to Causality

𝑞 𝑌5 ? 𝑌0 ≔ 𝑦

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For a particular realization of all RVs, What would have happened (to downstream variables) If 𝑁 had taken a different value? (keeping all other sources of randomness fixed) = 0 = 1 = 0 = 1 = 1 = 1 = 0 1 1 1 We can now differentiate: 𝑁 ⊕ 𝑎 is CCI’d by 𝑁, but 𝑎 is not!

Counterfactual Causal Influence (of 𝑁)

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= 0 = 1 = 0 = 1 = 1 = 1 = 0 1 1 1 We can now differentiate: 𝑁 ⊕ 𝑎 is CCI’d by 𝑁, but 𝑎 is not!

Counterfactual Causal Influence (of 𝑁)

Defining info flow using CCI:

  • 𝑁 may be constant over some
  • f its values 𝑛 ∈ ℳ

e.g. 𝑌 𝐹𝑢 = 𝕁 𝑁 ≥ 0

  • 𝑁 may be constant over all 𝑛

for some values of 𝑨 ∈ 𝒶 e.g. 𝑌 𝐹𝑢 = 𝑁 ⋅ 𝑎

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Definition [𝑁-Counterfactual Causal Influence]: We say that an edge 𝐹𝑢 is counterfactually causally influenced by 𝑁 if ∃ 𝑛, 𝑛′ ∈ ℳ and 𝑥 ∈ 𝒳 s.t. 𝑌 𝐹𝑢

𝑛,𝑥 ≠ 𝑌 𝐹𝑢 𝑛′,𝑥

𝑁-information flow 𝑁-counterfactual causal influence

Defining Info Flow using 𝑁-CCI

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Theorem [𝑁-CCI has no orphans]: If an outgoing edge 𝐹𝑢 of a node 𝑊

𝑢 is 𝑁-CCI’d, then there exists some

incoming edge 𝐹𝑢−1 of 𝑊

𝑢 that is also 𝑁-CCI’d.

Theorem [𝑁-CCI guarantees info paths]: If the transmissions of an “output” node 𝑊

𝑢 𝑝𝑞 depend on 𝑁, then there

is a path from 𝑊

𝑗𝑞 to 𝑊 𝑢 𝑝𝑞 such that every edge of this path is 𝑁-CCI’d.

Definition [𝑁-Counterfactual Causal Influence]: We say that an edge 𝐹𝑢 is counterfactually causally influenced by 𝑁 if ∃ 𝑛, 𝑛′ ∈ ℳ and 𝑥 ∈ 𝒳 s.t. 𝑌 𝑛,𝑥 𝐹𝑢 ≠ 𝑌 𝑛′,𝑥 𝐹𝑢

Main Results

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How else can we define Information Flow?

  • 1. Why study information flow?
  • 2. ISIT’19 recap: Defining information flow
  • Shortcoming: “𝑁-information Orphans”
  • Potential Fix based on Pruning
  • 3. Information Flow from a Causality Perspective
  • A counterexample to Pruning
  • Intro to Causality and Counterfactual Causal Influence
  • 4. CCI is the intuitive definition, but not observational
  • An Alternative Observational Definition
  • A Comparison of Definitions
  • 5. Conclusion
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CCI is not an Observational Measure

𝑞 𝑛, 𝑨, 𝑧 = 1 4 𝕁 𝑧 = 𝑛 ⊕ 𝑨 = 1 4 𝕁 𝑧 ⊕ 𝑨 = 𝑛

(using 𝑍 for 𝑁 ⊕ 𝑎)

Joint distribution is symmetric in 𝑁 ⊕ 𝑎 and 𝑎!

𝑁, 𝑎 ~ Ber(1/2) 𝑁 ⊥ 𝑎

  • Estimating CCI requires (intrinsic)

variables to be held fixed while 𝑁 is changed

  • Minimally requires the capability

to make interventions

  • Sometimes may be impossible to

estimate We would like observational measures. But…

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Can we find an alternative definition which does not suffer from the pruning counterexample? Definition [Modified 𝑁-Information Flow]: We say that an edge 𝐹𝑢 has modified 𝑁-information flow if ∃ ℰ𝑢

′ = 𝐹𝑢 𝑗 ⊆ ℰ𝑢 and functions 𝑔 𝑗 s.t. 𝐽 𝑁; 𝑌 𝐹𝑢 | {𝑔 𝑗 ∘ 𝑌 𝐹𝑢 𝑗 } > 0.

An alternative Observational Definition

𝑁

𝐵0 𝐶0 𝐵1 𝐶1 𝐵2 𝐶2

𝑊

𝑗𝑞

𝑊

𝑢 𝑝𝑞

𝐵3 𝐶3

𝑁 𝑁 𝑁 ⊕ 𝑎 [𝑁, 𝑎] 𝑁 ⊕ 𝑎 𝑁 𝑎 𝑎 𝑎 = 𝐽 𝑁; 𝑁 ⊕ 𝑎 𝑎 > 0 ∃ 𝑔: 𝑁, 𝑎 ↦ 𝑎 s.t. 𝐽 𝑁; 𝑁 ⊕ 𝑎 𝑔 𝑁, 𝑎

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All definitions have their shortcomings: 1. 𝑁-CCI identifies 𝑁 ⊕ 𝑎2 (but it cannot be decoded) 2. Modified 𝑁-info flow identifies 𝑎1 (but it cannot help decode 𝑁) 3. 𝑁-CCI ⇔ 𝑁-info flow 𝑁-CCI ⇔ Modified 𝑁-info flow 4. Modified 𝑁-info flow ⇒ 𝑁-info flow 𝑁-info flow 𝑁-CCI Modified 𝑁-info flow

Comparing the three definitions

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Summary and ongoing work

Information Flow & PID

(Venkatesh, Dutta and Grover, ISIT 2019; IEEE Trans. IT 2020, to appear)

Neuroengineering: Fundamental Limits and Algorithms

(Experimental validation: Robinson, Venkatesh et al., Sci. Rep. 2017) (Fundamental limits: Venkatesh and Grover, ISIT 2017) (with Improvements by Xu, Coleman in this ISIT!) (Theory: Grover and Venkatesh, Proc. IEEE 2016) (Clinical use case: Chaman Zar, …, Venkatesh et al., IEEE Trans. BME 2018)

AI and Fairness

(Dutta, Venkatesh, Mardziel, Datta and Grover, AAAI 2020)