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The computational brain (or “why studying the brain with math is cool”) Jonathan Pillow Princeton Neuroscience Institute & Psychology. +''+'+++' Math Tools for Neuroscience (NEU 314) Spring 2016

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Page 1: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the

The computational brain(or “why studying the brain with math is cool”)

Jonathan PillowPrinceton Neuroscience Institute & Psychology.

The University of Texas Department of Mathematics &

The Saturday Morning Math Group Present:

Featuring Dr. Jonathan Pillow of The University of Texas at Austin

For more info: www.math.utexas.edu/smmg

Information, Bits, Coding, and the Brain!

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Math Tools for Neuroscience (NEU 314)Spring 2016

Page 2: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the

What is computational neuroscience?

2. Study how the brain behaves as a computer• Brain is a machine for processing information &

computing relevant outputs

• What algorithms / routines does it use?

1. Use of mathematical/computational tools to study the brain.

• Estimate biological properties from noisy data

• build models that replicate behavior of neurons

Page 3: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the

Mind-Brain Problem

What is the relationship of the mind to the brain?

Page 4: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the

The brain as a computer:“The brain computes! This is accepted as a truism by the majority of neuroscientists engaged in discovering the principles employed in the design and operation of nervous systems. What is meant here is that any brain takes the incoming sensory data, encodes them into various biophysical variables, such as the membrane potential or neuronal firing rates, and subsequently performs a very large number of ill-specified operations, frequently termed computations, on these variables to extract relevant features from the input. The outcome of some of these computations can be stored for later access and will, ultimately, control the motor output of the animal in appropriate ways.”

- Christof Koch, Biophysics of Computation

Page 5: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the

Short history of brain metaphors:

• hydraulic device (Descartes, 17th C.)• mill (Leibniz, 17th C.)• telegraph (Sherrington, early 20th C.)• telephone switchboard (20th C.)• digital computer (late 20th C.)

• quantum computer? (Penrose, 1989)

• convolutional neural network? (21st C.)

Page 6: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the

What does it mean to claim the brain is a computer?

Answer to: What level of description suffices to explain how the brain gives rise to the mind?

Some possible answers: 1) Idealist/non-reductivist: None! No description of the brain could ever account for the properties of the mind.

Page 7: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the

Answer to: What level of description suffices to explain how the brain gives rise to the mind?

Some possible answers: 2) Physicalist: The brain’s physical properties are essential to generating the mind. No simulation or computational model suffices:

“As the liver secretes bile, so the brain secretes consciousness.”

What does it mean to claim the brain is a computer?

Page 8: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the

Answer to: What level of description suffices to explain how the brain gives rise to the mind?

Some possible answers: 3) Functionalist: A computational description. The physical properties are only important as place holders for the logical operations of a formal algorithm.

What does it mean to claim the brain is a computer?

Page 9: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the

BrainSensoryInput

MotorOutput

• The physical parts of the brain are important only insofar as they represent steps in a formal calculation.

• Any physical device implementing the same formal system would have the same “mind properties” as a brain.

What does it mean to claim the brain is a computer?

Page 10: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the

BrainSensoryInput

MotorOutput

Claim: Most neuroscientists take it for granted that the brain is a computer. They are devoted to finding out which computer (i.e., what formal structure? what algorithms does the brain implement?).

What does it mean to claim the brain is a computer?

Page 11: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the

Next: What is (some of) the evidence that the brain is a computer?

Page 12: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the

Mathematical model of sensory neurons

photoreceptors

bipolar cells

retinal ganglion cells

the retina

detect light

output cells (send all visual information to the brain)

to brain!

Page 13: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the

Mathematical model of sensory neurons

photoreceptors

bipolar cells

retinal ganglion cells

Difference of light in “center” and light in the “surround”

+ --

the retina

what mathematical operation?

Page 14: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the

Mathematical model of sensory neurons

photoreceptors

bipolar cells

retinal ganglion cells

Difference of light in “center” and light in the “surround”

+ --

stimulus

lots of spikes!

Page 15: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the

Mathematical model of sensory neurons

photoreceptors

bipolar cells

retinal ganglion cells

Difference of light in “center” and light in the “surround”

+ --

stimulus

few spikes

Page 16: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the

Mathematical model of sensory neurons

photoreceptors

bipolar cells

retinal ganglion cells

Difference of light in “center” and light in the “surround”

+ --

stimulus

more spikes

Page 17: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the

Mach Bands

Each stripe has constant luminance

Then why does it look like there’s a gradient?

Page 18: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the

Mach Bands

Each stripe has constant luminance

Then why does it look like there’s a gradient?

- + - Cell on right edge

- + -Cell on left edge

Page 19: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the

Lightness Illusion

Page 20: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the

Hermann illusion

Page 21: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the
Page 22: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the

Comparison patch

Page 23: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the

Comparison patch

Page 24: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the

Color Computations

Beau Lotto

Page 25: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the

Color Computations

Beau Lotto

Page 26: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the

sometimes this can fool 2/3 of the population (and sow division and hostility across the internet!)

Page 27: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the

Turns out: percept depends on statistical inferences brain makes about the light source!

Page 28: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the
Page 29: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the
Page 30: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the

color after-images

• neurons adjust their response properties after prolonged exposure to an image

• we can compute (and predict) these changes!• red —> green after-image • blue —> yellow after-image • black —> white after-image

Page 31: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the

Neural prostheses: Neurons can be replaced by other entities (silicon chips) that have different physical structure but carry out the same (or similar) mathematical operations, allowing the organism to produce (“compute”) the same behavior.

Page 32: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the

Cochlear implants(using a “different computer” to encode auditory signals)

microphone

transmitter receiver

cochlea

electrode array

to brain

Page 33: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the

Direct neural control of movement Schwartz Lab (Pitt)

Page 34: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the

Direct neural control of movement Schwartz Lab (Pitt)

Page 35: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the

If we understand the mathematical operations carried out by different parts of the brain, we could (in theory) replace them with new parts that perform the same computations!

Interchangeability: replacing neurons with silicon

BrainSensory

InputMotorOutput

Page 36: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the

10 microns

There are about 10 billion cubes of this size in your brain!

Page 37: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the

And this is a great time to study neuroscience. Why?

• We are about to get incredible data.

• Computers are getting extremely fast.

• Advances in statistical/mathematical techniques are allowing us to gain a deep understanding of neural data and neural information processing capabilities

Page 38: The Saturday Morning Math Group The …pillowlab.princeton.edu/teaching/mathtools16/slides/lec...The brain as a computer: “The brain computes! This is accepted as a truism by the

some of the kinds of math involved

• linear algebra

• probability & statistics

• dynamical systems / differential equations(including chaos theory)

• signal processing

• information theory / coding theory