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Introduction to Neuroscience: Systems Neuroscience Central visual processes Rafi Malach Department of Neurobiology

Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

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Page 1: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Introduction to Neuroscience: Systems Neuroscience

Central visual processes

Rafi Malach

Department of Neurobiology

Page 2: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

What is the function of the visual system?Creating an adaptive model of the environment

Page 3: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

The Visual Pathway

Page 4: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Low pass firing rate= BOLD fMRI

The Methods

Mukamel et al.

Firing rate

Functional Magnetic Resonance Imaging105-106 neurons

Hemodynamic Signal

Non InvasiveWhole brain coverage3X3 mm in 3T scanner

Page 5: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Micro-electrodes1-10 neurons

Firing rate

Contacts104-106 neurons

Broad band Gamma Power

Firing rate= amplitude of fast (Gamma) fluctuations

Electrophysiology

Nir et al.

Spikes

LFP

Spikes/Gamma

Page 6: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts
Page 7: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

retinotopy, visual field, contra- ipsi, fixation point, vertical meridianHorizontal meridian

Definitions:

Page 8: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

YHIVZHFNXR

Optic nerve, chiasm, tract and radiation

Flow of information form the eye to the brain

Page 9: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Atlas of human visual areas

Page 10: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Unfolded atlas of human visual areas

Page 11: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

The lay-out of early visual areas

Page 12: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Visual selectivity of single neurons:

a. What is its function in visionb. How is it generated

Visual Patterns

...

Firing Rate

1 2 3…

Page 13: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

“Tuning curve”

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Firing Rate

Visual Parameter(e.g. retinal position)

Sharply or narrowly tuned, selective

Page 14: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Invariance: lack of sensitivity to changes in a visual property

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Firing Rate

Visual Parameter(e.g. retinal position)

Page 15: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

The properties of single neurons in area V1

The combinatorial explosion problem:The number of possible visual patterns is ultra-vast

Page 16: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

The properties of single neurons in area V1

David Hubel and Torsten Wiesel

Page 17: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

The properties of single neurons in area V1

Page 18: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Receptive field of a visual neuron in area V1

Page 19: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Columns: common vertical specialization

Page 20: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

What is the function of the oriented line detectors?

Page 21: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Perceptual saliency of line detection

Page 22: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

How is orientation selectivity generated?

Page 23: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Receptive field of a “Simple” cell in area V1

Stimulus selectivity of receptive fields

LGN

“Bar”

“Edge”

Page 24: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

The simple cell model

Convergence, threshold, synchronyAn “and” function

Page 25: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

The cortex is organized in layers

Inputs from

Lower Centers

Outputs to

higher centers

Outputs to other

systems

Page 26: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Ocular Dominance Columns

Page 27: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Monkey Ocular Dominance Columns- top view

Page 28: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

What is the function of the ocular selectivity?

Page 29: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Orientation and ocular columns- the “hyper –column”

Page 30: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Orientation columns- top view

Orientation vs. OD

Page 31: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Lateral Connectivity: massive, local, reciprocal interactions

0.2 mm

Page 32: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Lateral Connectivity connects similar-function columns

Page 33: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

What is the function of the lateral connectivity

++

+

+

Positive feedback:“Ignition”- non linear response curves“attractors”- Completion effects“Recurrence”- Short term memory

Page 34: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Demonstration of non-linear “Ignition-like” behavior

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+

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Page 35: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts
Page 36: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Demonstration of non-linear “Ignition-like” behavior

++

+

+

Page 37: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Demonstration of completion effects

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+

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Page 38: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Large scale organizational principles of V1

Retinotopy

Mmagnification factor- mm cortex/visual angle

Page 39: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Polar Retinotopic organization of visual cortex

HM

VM

WORLD CORTEX

HM VM

fovea

periphery

00 900TETA

R

Page 40: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Center

Periph.

Center-Periphery organization

Page 41: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

What happens down-stream?

Low

High

Page 42: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Complex Receptive field of a visual neuron in area V1

Page 43: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Complex Receptive field of a visual neuron in area V1How is it generated?

Page 44: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Complex Receptive field of a visual neuron in area V1What is its function?

Position invariance

Page 45: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Fusiform “Face” Area

Page 46: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Parahippocampal “Place” Area

Page 47: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Complex templates at the top of the VENTRAL stream

“Face” neurons

Page 48: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Face “Patches” are built of “face-neurons”

D. Tsao

Page 49: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Electrical stimulation of the fusiform face area

Page 50: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Columnar organization of complex shapes in high order Ventral Stream

Page 51: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Large scale principles: Category organization

Page 52: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Growing “abstraction” along the visual hierarchy

Increased complexity

Increased RF size

Page 53: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

A toy model of integration along the cortical hierarchy

How can we advance beyond the toy model approach?

Page 54: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Artificial networks now out-perform human visual recognition capabilities

Deep Convolutional Neural Network

Yamin and DiCarlo, 2016Guculu and Van Gerven, 2015

Page 55: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

How can Deep networks compared to brain networks?

Page 56: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Exemplar selectivity can be converted to distance between pictures

Similar activation patterns

Short distance (ED=0.77)

Dissimilar activation patterns

Long distance (ED=1.28)

Page 57: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Comparing neural activation patterns driven by

different faces to DCNN activation patterns

Similar activation patterns

Different activation patterns

Neural distances matrix

Page 58: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Hierarchy-based correlation of Human visual areas and Deep Network

Page 59: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Hierarchy-based correlation of Human visual areas and Deep Network

Page 60: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Large scale specialization

Page 61: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Two streams in the visual system

Page 62: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Content-Selectivity in human visual cortex

Gestalttemplates

IT/FFA

Motion(MT/V5)

Local elementsV1

Dorsal-stream(Action)

Ventral-stream(Recognition)

Page 63: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Direction selectivity in area MT- Dorsal stream

Page 64: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts
Page 65: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Columnar organization for direction and orientation in area MT

Page 66: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts
Page 67: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Electrical stimulation of MT direction column- shifts perceptual judgment

Page 68: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Motion blindness following dorsal stream lesion

Page 69: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Action specialization in dorsal stream cortex

Page 70: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Shmuelof and Zohary

Action specialization in dorsal stream cortex

Page 71: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Massive “Top-Down” connections along the visual hierarchy

Page 72: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Attention mediated by top-down connections

Moran and Desimone

Page 73: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Top down attention in the human visual system

Page 74: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Spatial (“spot-light”)attention in early visual system

Page 75: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Template –selective attention in high order, ventral stream

Page 76: Introduction to Neuroscience: Systems Neuroscience Central … · 2018-12-01 · Whole brain coverage 3X3 mm in 3T scanner. Micro-electrodes 1-10 neurons Firing rate Contacts

Spatial attention is mediated via the right dorsal stream

Visual Neglect following right parietal lobe lesion