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Fundamentals of area V1 Part 1: Receptive fields and maps Matteo Carandini Smith-Kettlewell Eye Research Institute www.ski.org/Carandini

Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

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Page 1: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Fundamentals of area V1Part 1: Receptive fields and maps

Matteo CarandiniSmith-Kettlewell Eye Research Institute

www.ski.org/Carandini

Page 2: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Dowling, 1987 (Fig 1.2)

Page 3: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Receptive fields in area V1

Hubel and Wiesel, circa 1969 (from Nicholls et al., 1992)

Page 4: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Responses of a V1 neuron

Hubel and Wiesel, 1959

Page 5: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Artificial early visual system

www.ini.unizh.ch/~tobi/friend/chip/

Delbruck & Liu, 2004

Page 6: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Responses of artificial simple cell

Receptive fieldTuning curve

Delbruck & Liu, 2004

Page 7: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Receptive field of a V1 simple cell

DeAngelis, Ohzawa & Freeman, 1995

Page 8: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Receptive field of a V1 simple cell

DeAngelis, Ohzawa & Freeman, 1995

Page 9: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Selectivity for orientation

Page 10: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Stimuli and receptive fields in space-time

Adelson & Bergen, 1985 Carandini, Heeger & Movshon , 1999

Page 11: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Separability and direction selectivity

Separable,not direction selective

Inseparable,direction selective

Page 12: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Space-time receptive fields

DeAngelis, Ohzawa & Freeman, 1995

Page 13: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

V1 simple cell, separable

www.bpe.es.osaka-u.ac.jp/ohzawa-lab/ Ohzawa, DeAngelis & Freeman, 1996

Page 14: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

V1 simple cell, inseparable

www.bpe.es.osaka-u.ac.jp/ohzawa-lab/ Ohzawa, DeAngelis & Freeman, 1996

Page 15: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

How are V1 receptive fields obtained?

DeAngelis, Ohzawa & Freeman, 1995

Page 16: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Assembly of receptive fieldsin the artificial visual system

ON center ganglion cell(Vm: 2, R: 3)

OFF center ganglion cell(R: 4)

ODD simple cell(Vm: 5, R: 7)

Photoreceptor (1)

Horizontal cell (average photoreceptor value)

EVEN simple cell(Vm: 6, R: 8)

www.ini.unizh.ch/~tobi/friend/chip/ Delbruck & Liu, 2004

Page 17: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

H & W’s feedforward model of simple cells

Visual field Brain

LGNcells

Simplecell

Hubel & Wiesel, 1963

Page 18: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Receptive field of a V1 complex cell

ON

DeAngelis, Ohzawa & Freeman, 1995

Page 19: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Receptive field of a V1 complex cell

OFF

DeAngelis, Ohzawa & Freeman, 1995

Page 20: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

H & W’s feedforward model of complex cells

Visual field Brain

Simplecells

Complexcell

Hubel & Wiesel, 1963

Page 21: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Simple cell responses to a drifting grating

Page 22: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Responses to a drifting grating

Carandini & Ferster, 2000

Page 23: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Responses to a drifting grating

Carandini & Ferster, 2000

Page 24: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Dichotomy of simple and complex cellsComplex Simple

Cat

Monkey

F1/F0Skottun, DeValois, Grosof, Movshon, Albrecht & Bonds , 1991

Page 25: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Responses to a drifting grating

Carandini & Ferster, 2000

Page 26: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Dichotomy is created by threshold

Priebe, Mechler, Carandini & Ferster, 2004

Page 27: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Feedforward model of complex cells

C

Page 28: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Complex cells as networked simple cells

Low gain

Chance, Nelson & Abbott, 1999

Page 29: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Complex cells as networked simple cells

C

C

CHigh gain

C

Chance, Nelson & Abbott, 1999

Page 30: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Complex cells as networked simple cells

Gain = 1

Gain = 5

Gain = 20

Simple

Intermediate

Complex

Chance, Nelson & Abbott, 1999

Page 31: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Maps

Page 32: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Cortical representationmeasured with 2-deoxy-glucose

Tootell et al. (1988)

Page 33: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Complex-log model

Schwartz et al. (1988)Frederick and Schwartz (1990)

Page 34: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

If your eyes see this…

Schwartz et al. (1988)Frederick and Schwartz (1990)

Page 35: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

…your brain maps it to this

Schwartz et al. (1988)Frederick and Schwartz (1990)

Page 36: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Retinotopy is very precise

Adams & Horton (2003)

Page 37: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Map of ocular dominancemeasured with radioactive proline

LeVay, Hubel and Wiesel (1975)in Nicholls et al. (1992)

Page 38: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Optical imaging

•Intrinsic signals•Voltage-sensitive dye

Page 39: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Map of ocular dominancemeasured with optical imaging

Bonhoeffer & Grinvald (1991)in Nicholls et al. (1992)

Page 40: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Map of orientation preferencemeasured with optical imaging

Bonhoeffer and Grinvald (1991)in Nicholls et al. (1992)

Page 41: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Relationship between ocular dominance and orientation preference

Obermayer & Blasdel (1993)

Page 42: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Connections from LGN may constrainthe map of orientation preference

Page 43: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Preliminary evidence from tree shrew V1

Mooser, Bosking & Fitzpatrick, Soc Neurosci Abs 2001

Page 44: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

The retinal mosaic may determine orientation preference

Retinal mosaic, X cells

Ringach, 2004

Page 45: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

The retinal mosaic may constrainthe map of orientation preference

Retinal mosaic, X cells

Ringach, 2004

Page 46: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Summary

• V1 receptive fields in space-time• Wiring of simple cells and complex cells• Maps of selectivity• Constraints on orientation map

Page 47: Fundamentals of area V1 Part 1: Receptive fields and maps · 2004-07-09 · Tuning curve Receptive field Delbruck & Liu, 2004. Receptive field of a V1 simple cell DeAngelis, Ohzawa

Power law creates dichotomy

Priebe, Mechler, Carandini & Ferster, 2004