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An Introduction to the Visual System

An Introduction to the Visual System

Second edition

Martin J. Tove´e

Newcastle University

CAMBRIDGE UNIVERSITY PRESS

Cambridge, New York, Melbourne, Madrid, Cape Town, Singapore, São Paulo

Cambridge University Press

The Edinburgh Building, Cambridge CB2 8RU, UK

Published in the United States of America by Cambridge University Press, New York

www.cambridge.org

Information on this title: www.cambridge.org/9780521883191

© M. J. Tovee 2008

This publication is in copyright. Subject to statutory exception and to the provision of relevant collective licensing agreements, no reproduction of any part may take place without the written permission of Cambridge University Press.

First published in print format 2008

ISBN-13

978-0-511-41393-3

eBook (EBL)

ISBN-13

978-0-521-88319-1

hardback

ISBN-13

978-0-521-70964-4

paperback

Cambridge University Press has no responsibility for the persistence or accuracy of urls for external or third-party internet websites referred to in this publication, and does not guarantee that any content on such websites is, or will remain, accurate or appropriate.

This book is dedicated to my wife Esther, and

to our children Charlotte and James.

Contents

1

 

Introduction

1

 

 

A user’s guide?

1

 

 

Brain organisation

2

 

 

Why is the cerebral cortex a sheet?

4

 

 

Cortical origami

6

 

 

Does connectivity predict intelligence?

7

 

 

Analysis techniques: mapping the brain

8

 

 

Structural imaging

8

 

 

Functional imaging techniques: PET and fMRI

10

 

 

What is the relationship between blood flow

 

 

 

and neural activity?

12

 

 

The resolution problem

13

 

 

Measuring brain activity in real time: MEG and EEG

14

 

 

Transcranial magnetic stimulation (TMS)

15

 

 

Summary of key points

16

 

 

 

2

 

The eye and forming the image

18

 

 

 

What is the eye for?

18

 

 

Light

18

 

 

The structure of the eye

19

 

 

Focusing the image

25

 

 

The development of myopia

26

 

 

Clouding of the lens (cataracts)

28

 

 

Photoreceptors

28

 

 

Transduction

30

 

 

The calcium feedback mechanism

31

 

 

Signal efficiency

32

 

 

The centre-surround organisation of the retina

33

 

 

Light adaptation

36

 

 

Duplicity theory of vision

37

 

 

Sensitivity, acuity and neural wiring

40

 

 

Summary of key points

41

 

 

 

 

3

 

Retinal colour vision

44

 

 

Why do we need more than one cone pigment?

44

 

 

Trichromacy

44

 

 

The genetics of visual pigments

47

 

 

The blue cone pigment

53

 

 

Rhodopsin and retinitis pigmentosa

54

viii C O N T E N T S

 

 

Better colour vision in women?

55

 

 

Three pigments in normal human colour vision?

56

 

 

The evolution of primate colour vision

59

 

 

What is trichromacy for?

59

 

 

Summary of key points

60

 

 

 

 

4

 

The organisation of the visual system

62

 

 

 

Making a complex process seem simple

62

 

 

The retina

63

 

 

The lateral geniculate nucleus (LGN)

63

 

 

The primary visual cortex (VI)

64

 

 

Visual area 2 (V2)

67

 

 

Visual area 4 (V4)

68

 

 

Visual areas 3 (V3) and 5 (V5)

69

 

 

The koniocellular pathway

69

 

 

The functional organisation

70

 

 

Perception vs. action

71

 

 

Blindsight

73

 

 

Summary of key points

76

 

 

 

 

5 Primary visual cortex

78

 

 

The visual equivalent of a sorting office?

78

 

 

Segregation of layer 4 inputs

79

 

 

Cortical receptive fields

79

 

 

Spatial frequency

81

 

 

Texture

82

 

 

Direction selectivity

82

 

 

Colour

84

 

 

Modular organisation

84

 

 

Summary of key points

87

 

 

 

 

6

 

Visual development: an activity-dependent process

89

 

 

Variations on a theme

89

 

 

Monocular or binocular deprivation

91

 

 

Image misalignment and binocularity

93

 

 

Image misalignment in humans

94

 

 

Selective rearing: manipulating the environment

96

 

 

Impoverished visual input in humans

98

 

 

The critical period

98

 

 

What we see, shapes how we see it

99

 

 

Summary of key points

99

C ON T EN T S ix

7

 

Colour constancy

101

 

 

The colour constancy problem

101

 

 

The Land Mondrian experiments

102

 

 

Reflectance and lightness: the search for constancy

 

 

 

 

in a changing world

103

 

 

The biological basis of colour constancy

105

 

 

Colour constancy and the human brain

106

 

 

Summary of key points

108

 

 

 

 

8

Object perception and recognition

109

 

 

From retinal image to cortical representation

109

 

 

Early visual processing

109

 

 

A visual alphabet?

112

 

 

Complex objects in 3-D: face cells

118

 

 

Functional divisions of face cells: identity, expression

 

 

 

 

and direction of gaze

120

 

 

The grandmother cell?

121

 

 

Are face cells special?

122

 

 

Visual attention and working memory

126

 

 

Fine-tuning memory

129

 

 

A clinical application?

130

 

 

Visual imagery and long-term visual memory

131

 

 

Summary of key points

132

 

 

 

 

9

Face recognition and interpretation

133

 

 

What are faces for?

133

 

 

Face recognition

133

 

 

Laterality and face recognition

136

 

 

How specialised is the neural substrate of face

 

 

 

 

recognition?

138

 

 

The amygdala and fear

139

 

 

The frontal cortex and social interaction

143

 

 

Faces as a social semaphore

144

 

 

Summary of key points

145

 

 

 

 

 

10

 

Motion perception

147

 

 

 

 

The illusion of continuity

147

 

 

 

Saccades

148

 

 

 

Suppression of perception during saccades

150

 

 

 

What happens if you don’t have saccades?

151

 

 

 

How to stabilise the visual world

152

 

 

 

Navigating through the world: go with the flow?

153

x C O N T E N T S

 

Going against the flow?

155

 

The neural basis of motion detection

156

 

Human V5

161

 

Summary of key points

163

 

 

 

11

Brain and space

164

 

The final frontier

164

 

Oculomotor cues

164

 

Interposition

165

 

Relative size

166

 

Perspective

166

 

Motion parallax

168

 

Stereopsis

168

 

The neural basis of three-dimensional

 

 

space representation

169

 

The problem of visual neglect

170

 

The neural basis of neglect

172

 

Summary of key points

174

 

 

 

12

What is perception?

175

 

Putting it all together

175

 

Neuronal oscillations

175

 

How else to solve the problem

178

 

What is perception?

180

 

Change blindness

180

 

Perceptual rivalry

182

 

The illusion of perception

185

 

Summary of key points

185

References

187

Index

210

The colour plates are to be found between p. 88 and p. 89.