Visual Development - 3rd Edition

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Nigel W. Daw
Branford, CT, USA

ISBN 978-1-4614-9058-6 ISBN 978-1-4614-9059-3 (eBook)

DOI 10.1007/978-1-4614-9059-3
Springer New York Heidelberg Dordrecht London

Library of Congress Control Number: 2013949433

1st edition: © Plenum Press, New York 1995

2nd edition: © Springer Science + Business Media, Inc 2006
© Springer Science+Business Media New York 2014
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Preface

Research in the area of visual development has become a multidisciplinary

affair. Students who acquire an interest in the field therefore need to under-
stand several different aspects of the subject. The development of acuity mea-
sured by psychophysicists is the concern of optometrists and ophthalmologists,
and depends on changes in the anatomy of the retina and the physiology of
cells in the visual pathway. Scientists working on the cellular, molecular, and
biochemical mechanisms lean on anatomical studies, physiology, and psy-
chophysics in designing and interpreting their experiments. Indeed, the labo-
ratories of the leading scientists working on the subject now all use a large
variety of techniques in their studies.
Because the study of visual development is pursued by workers in many
disciplines, from medicine to basic science, I have tried to write this book at
a level at which it can be understood by a variety of students: graduate stu-
dents in neurobiology and psychology, as well as optometry students and
ophthalmology residents. The text assumes some knowledge of basic terms
such as acuity, but a glossary is provided should the reader find some words
that are unfamiliar. The emphasis is on facts and conclusions, rather than on
methods and procedures. Many details are left out.
However, I hope that the experts will also read the book. The subject has
become so wide-ranging that not many people have the time to read literature in
all aspects of it. The book is also intended for experts in one area to get a grasp of
the basics of the subject in other disciplines that are not their primary discipline.
To write a book covering such a wide variety of disciplines, I have had to
simplify. The book does not go into controversies in detail. Instead, it pro-
vides my summary of what seems to me to be the best evidence. Not every-
body will agree with my synthesis. Some experts will read it and be outraged
at some of my statements. However, my outrageous statements were intended
to be constructive: I hope that they will provoke thought and point the way to
more experiments that will carry the field forward.
I am grateful to Colin Blakemore for inviting me to write this book. The
process has been an educational one for me, and led to a number of insights
which may have been apparent to others, but not to me. A number of friends
and colleagues have helped me in the preparation of the book. Grace Gray
in particular read the entire text twice, and improved it throughout. Robert
Hess went through the whole section on Visual Deprivation, and made

v
vi Preface

many valuable comments. John Lisman did the same for the section on
Mechanisms. Janette Atkinson, Marty Banks, Oliver Braddick, Jan Naegele,
Pasko Rakic, and Josh Wallman read individual chapters in their area of
expertise and made many corrections and improvements. Several of my col-
leagues in the Department of Ophthalmology—Ethan Cohen, Jonathan
Kirsch, Thomas Hughes, Colin Barnstable, Silvia Reid and Helen Flavin—
gave comments on various portions of the text, and Marc Weitzman read
two whole sections. I would like to thank them all. However, I did not adopt
all of their suggestions, and the errors and omissions are mine. I would also
like to thank Janet Hescock and Bob Brown for help in the preparation of
the text and figures, together with support from the Core Grant to Yale
University from the National Eye Institute.

Branford, CT, USA Nigel W. Daw

Preface to the Third Edition

It is almost 20 years since the first edition of this book was written, and con-
siderable research has been done in that time. The development of vernier
acuity and contour discrimination have been more carefully defined. Genes
that play a role in myopia have been localized. New techniques using Gabor
patches in various configurations have enabled scientists to define amblyopia
in more careful terms, particularly the concept of spatial uncertainty, and
whether it is due to undersampling or distorted sampling. Amblyopia was
never simply a matter of a deficit in acuity, but it has taken careful experi-
ments to show exactly what it is beyond that.
Ten years ago, the molecules that govern the crossing of the optic nerve
fibers in the chiasm, and which project to the contralateral side, and which to
the ipsilateral side were completely unknown. So were the molecules that
govern the topography of the projections within the visual system. Today, we
know some molecules involved in both these developmental events, as well as
some that mark the boundaries of the visual cortex.
The technique of optical imaging of the visual cortex has enabled scien-
tists to visualize the ocular dominance and orientation columns. Scientists
can now use this technique to study the development of these columns, and
the effect of various forms of visual deprivation on them, in a way that was
not possible with single unit recordings.
It has also become increasingly apparent that there are many critical peri-
ods in the development of the visual system. The critical period for the effect
of a particular form of visual deprivation in many cases lasts longer than the
period of development of the property affected, and the period during which
recovery can be obtained lasts longer still. Moreover, there are different criti-
cal periods for different properties. Properties handled at a higher level of the
system have a later critical period. In addition, the critical period can be
affected by the previous visual history of the animal, and by the technique
used to evaluate it. For all these reasons, the chapter on critical periods has
become much more involved.
Twenty years ago, quite a lot was known about mechanisms of long-term
potentiation and long-term depression, and not much about plasticity in the
visual cortex resulting from monocular deprivation. Today, a considerable
amount is known about mechanisms of ocular dominance plasticity, and what
is known about LTP and LTD does not add a great deal, so the chapter on the
latter has been eliminated.

vii
viii Preface to the Third Edition

The subject of the effect of out-of-focus images on the size of the eyeball has
also been a very active area of research. We still do not know how the signal
gets from the neural retina to the choroid and sclera, but there are a few mole-
cules known that increase in response to plus lenses and decrease in response to
minus lenses, or vice versa, that may be candidates for the signal.
Moreover, the best treatment for amblyopia has become much better
known, as therapists concentrate on properties other than acuity. Basic scien-
tists have quantified the improvements that can be made by not patching the
amblyopic eye all of the time, and working with both eyes to improve binocu-
lar vision at the same time as the acuity in the amblyopic eye. Use of percep-
tual learning and video games has helped by increasing activity and attention
as the therapy is done. Many of the principles have been employed by pediat-
ric vision therapists for some time, but the publicity generated by “Stereo
Sue” and others has helped to broadcast them.
For all these reasons, it is high time that this book should be revised. The
aim of the book is the same as it was in the first edition—to provide a short
summary of findings in the field that can be used by ophthalmology residents,
optometry students, graduate students in neurobiology and psychology,
senior undergraduates, and, since the field is so diverse, for experts to read
chapters outside their area of expertise.
For this third edition, there is a Web site, www.visual-development.net,
with links to various videos that illustrate points and procedures discussed in
the book. See also the list of useful videos after the table of contents.
I am most grateful to Paul Harris for reading the whole book and providing
a number of suggestions for clarification, as well as for professional guidance
on the new chapter on “Treatment of Amblyopia.” I am also very grateful to
Terri Lewis, Len White, Mike Crair, Eileen Birch, Dennis Levi, Donald
Mitchell, John Lisman, Takao Hensch, and David Troilo, who provided com-
ments on individual chapters and hopefully caught most of my errors, but I am
sure that some remain. I also thank Simina Calin, my editor at Springer, for
all her help and guidance.
Contents

1 Introduction .................................................................................. 1
References ...................................................................................... 5
2 Functional Organization of the Visual System .......................... 7
General Anatomical Organization ................................................. 8
Function in the Retina .................................................................... 11
Function in the Lateral Geniculate Nucleus................................... 14
Function in the Visual Cortex ........................................................ 16
The Columnar Organization of Cortex ...................................... 17
Parallel Processing Within the Visual System ............................... 19
Hierarchical Processing Within the Visual System........................ 19
Higher Visual Areas ................................................................... 22
Summary ........................................................................................ 22
References ...................................................................................... 23

Part I Development of the Visual System

3 Development of Visual Capabilities............................................ 27

Methods for Studying Infant Vision .............................................. 28
What Can Be Seen ......................................................................... 30
Faces .......................................................................................... 30
Biological Motion and Optic Flow ............................................ 30
Contour Integration, Illusory Contours,
and Continuity Behind Other Objects ........................................ 31
Depth Perception........................................................................ 32
Fusion and Orthophoria ............................................................. 33
Color .......................................................................................... 34
Vision in the Periphery of the Visual Field ................................ 34
What Can Be Measured ................................................................. 35
Acuity......................................................................................... 35
Contrast Sensitivity .................................................................... 36
Vernier Acuity ............................................................................ 39
Stereo ......................................................................................... 40
Prestereopsis and Poststereopsis ................................................ 42
The Prestereoptic and Poststereoptic Periods ............................ 43
Orientation, Direction, Movement ............................................. 43
Adaptation .................................................................................. 46

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x Contents

Development of Eye Movements ................................................... 46

Fixation and Refixation .............................................................. 46
Saccades ..................................................................................... 46
Smooth Pursuit ........................................................................... 48
Vergence and Accommodation .................................................. 48
Optokinetic Nystagmus.............................................................. 49
Summary ........................................................................................ 50
References ...................................................................................... 50
4 Anatomical Development of the Visual System ......................... 55
Development of the Retina and Its Projections .............................. 56
Crossing in the Optic Chiasm .................................................... 57
Errors at the Chiasm in Albino Animals .................................... 59
Development of the Lateral Geniculate Nucleus ........................... 60
Topography of Projections from Retina ......................................... 61
Cell Connections Within the Overall Topographic Projection... 64
Development of Visual Cortex and Projections to It ..................... 64
Development of Geniculocortical Projections ........................... 65
Formation of Layers in the Cortex ............................................. 65
Role of the Subplate Neurons .................................................... 66
Migration of GABA Cells .......................................................... 67
Development of Connections to and from Layers
in the Visual Cortex.................................................................... 68
Development of Clusters of Cells with Similar Properties ............ 69
Formation of Ocular Dominance Columns ................................ 70
Formation of Blobs and Pinwheels ............................................ 70
Development of Lateral Connections ........................................ 70
Other Events During Differentiation.............................................. 70
References ...................................................................................... 73
5 Development of Receptive Field Properties ............................... 77
Development in the Retina and Lateral Geniculate Nucleus ......... 79
Development in Primary Visual Cortex ......................................... 81
Other Parts of Cortex Concerned with Vision ............................... 87
Development in the Absence of Light and Activity....................... 88
References ...................................................................................... 90

Part II Amblyopia and the Effects of Visual Deprivation

6 Modifications to the Visual Input That Lead

to Nervous System Changes ........................................................ 95
Strabismus ...................................................................................... 96
Anisometropia ................................................................................ 100
Astigmatism ................................................................................... 100
Cataract and Corneal Opacities ...................................................... 102
Myopia ........................................................................................... 103
References ...................................................................................... 104
Contents xi

7 Known Physiological and Anatomical Changes That Result

from Optical and Motor Deficits................................................. 107
Monocular Deprivation .................................................................. 108
Binocular Deprivation and Congenital Blindness .......................... 112
Retinal Lesions............................................................................... 112
Orientation and Direction Deprivation........................................... 113
One Eye Out of Focus .................................................................... 114
Strabismus ...................................................................................... 116
References ...................................................................................... 120
8 What Is Amblyopia? .................................................................... 123
Amblyopia from Anisometropia .................................................... 124
Effect of Cataracts.......................................................................... 125
Amblyopia from Strabismus .......................................................... 126
Properties Affected in a Variety of Causes of Amblyopia ............. 128
Crowding.................................................................................... 130
Vernier Acuity ............................................................................ 132
Spatial Uncertainty..................................................................... 133
Components of the Loss in Acuity in Various Strabismics............ 137
Detailed Comparison of Groups of Amblyopes............................. 138
The Peripheral Field of View ......................................................... 139
A Rapid Test for Amblyopia .......................................................... 140
Eye Movements in Amblyopes ...................................................... 140
Fixation ...................................................................................... 140
Saccades ..................................................................................... 142
Reading ...................................................................................... 142
Summary ........................................................................................ 142
References ...................................................................................... 143
9 Critical Periods............................................................................. 147
General Principles from Experiments with Animals ..................... 148
The System Is Most Plastic Between Eye Opening
and Puberty ................................................................................ 148
More Severe Deprivations Have a Larger Effect ....................... 149
Higher Levels of the System Have a Later Critical Period ........ 149
Different Properties Have Different Critical Periods ................. 150
The Critical Period Depends on the Previous Visual
History of the Animal ................................................................ 152
Procedures Affect the Critical Period ........................................ 152
The Periods of Development, Disruption,
and Recovery May Be Different ................................................ 154
Critical Periods in Humans ............................................................ 155
Disruption of Acuity .................................................................. 155
Recovery from Disruption of Acuity ......................................... 157
Binocularity................................................................................ 158
Stereopsis ................................................................................... 159
Movement .................................................................................. 160
Plasticity and Learning in the Adult .......................................... 162
Summary ........................................................................................ 162
References ...................................................................................... 163
xii Contents

10 Treatment of Amblyopia .............................................................. 167

Optical and Motor Treatment ......................................................... 168
Patching.......................................................................................... 168
Equalizing the Monocular Skills in the Two Eyes ......................... 169
Binocularity and Fusion ................................................................. 170
Stereopsis ....................................................................................... 172
Eye Movements.............................................................................. 173
Activity During Therapy ................................................................ 174
Perceptual Learning ................................................................... 174
Video Games .............................................................................. 175
Pointers from Animal Experiments ............................................... 176
Sequencing of Treatment ............................................................... 177
Treatment of Expectations ............................................................. 177
Success Rates ................................................................................. 178
References ...................................................................................... 178

Part III Mechanisms of Plasticity

11 Concepts of Plasticity................................................................... 183

The Hebb Postulate ........................................................................ 184
How Electrical Activity Can Strengthen Some Synapses
and Weaken Others ........................................................................ 185
Feedback from the Postsynaptic Cell to the Presynaptic Terminals.... 186
Criteria for Critical Factors in the Critical Period ......................... 187
References ...................................................................................... 189
12 Plasticity in the Visual Cortex..................................................... 191
Electrical Activity .......................................................................... 192
Polarization of the Postsynaptic Neuron ........................................ 193
NMDA Receptors .......................................................................... 195
Metabotropic Glutamate Receptors ............................................... 197
GABA and Excitation-Inhibition Balance ..................................... 198
Brain-Derived Neurotrophic Factor (BDNF) and Insulin-Like
Growth Factor-1 (IGF-1), and Tumor Necrosis Factor α ............... 200
Calcium, α-Calcium/Calmodulin Kinase, and Calcineurin ........... 201
cAMP and Protein Kinase A .......................................................... 202
A Kinase Anchoring Protein at the Postsynaptic Density ............. 204
Other Protein Kinases ................................................................ 204
Cyclic AMP Response Binding Element (CREB) ......................... 204
Genes and Protein Synthesis .......................................................... 204
Structural Factors ........................................................................... 206
Modulatory Factors ........................................................................ 207
Nerve Growth Factor (NGF) ...................................................... 208
Immune System Molecules ............................................................ 208
Different Mechanisms for Different Aspects of Plasticity............. 209
Factors to Increase Plasticity ..................................................... 209
Summary ........................................................................................ 210
References ...................................................................................... 210
Contents xiii

13 Visually Induced Myopia and Emmetropization ...................... 217

Emmetropization ............................................................................ 217
Local Signals Go from the Retina to the Eyeball .......................... 218
Sensitive Period for Changes in Eyeball Growth ........................... 220
Changes in Both Choroid and Sclera ............................................. 221
Mechanisms for the Signals Going from the Retina
to the Eyeball ................................................................................. 221
Genes Associated with Myopia ..................................................... 225
Possible Treatments ....................................................................... 225
References ...................................................................................... 227

Glossary ................................................................................................ 231

Index ...................................................................................................... 241

List of Useful Videos

This book has a website associated with it—www.visual-development.net.

The videos listed below can all be accessed from the relevant paragraph in the
book, or from this website. As new videos become available, they will be
added to the website.

For Chap. 1
A discussion of Molyneux’s question is given by Dick Held at http://youtu.
be/EDRa-ESxmJY.

For Chap. 2
Recordings at various levels of the visual system by Clay Reid http://youtu.
be/mtPgW1ebxmE.

For Chap. 3
A demonstration of Biological Motion by Sverker Runesson at http://www.psyk.
uu.se/organisation/historia/?languageId=1 including motion demonstrations
in both 2 and 3 dimensions from the work of Professor Gunnar Johansson.
Development of acuity and face perception by David Martin http://youtu.be/
GK3ebhSmC4A.
Development of acuity, depth and color by Sepura Dosetareh http://youtu.be/
wsPIIC2b2wA.
A description of what babies see by Terri Lewis http://youtu.be/
UGqQnR5weTg.

For Chap. 4
A basic description of the development of the brain, focusing on hearing
rather than vision, is given by Joan Stiles at http://youtu.be/FugrcVhi2tg.
Differentiation of neural stem cells and development of cerebral cortex by
Pasko Rakic http://youtu.be/jeSh9t4aMa4.

For Chaps. 4 and 5

Comparison of organization of orientation columns in various species by Len
White www.visual-development.net.
xv
xvi List of Useful Videos

For Chaps. 5 and 7

Comparing the development of mouse and macaque visual systems and the effects
of visual deprivation by Michael Stryker http://youtu.be/W_MwoORWnlI.

For Chaps. 6 and 10

A description of amblyopia by Dominick Maino http://youtu.be/2tiV5vmHaEg.

For Chap. 8
A rapid test for amblyopia by Denis Pelli at http://psych.nyu.edu/pelli/
dualacuity/.

For Chaps. 9 and 12

Critical periods and mechanisms of plasticity by Takao Hensch http://youtu.
be/xZvqAHhiEL8.

For Chap. 10
A demonstration of the Brock Posture Board by Paul Harris at http://youtu.
be/CEKqNTV8Q3k.
A demonstration of the Brock String by lDoc 2008 at http://youtu.be/
EGlCVTdNqfw.
A demonstration of the Wayne Saccadic Fixator by visionforlifeworks at
http://youtu.be/K3UfrB_VNSw.
Various vision therapy exercises by tglass0000 at http://youtu.be/
MWKWNPdxpyk.
Video games, and how the two eyes are made to work together by Jessica
Bayliss http://youtu.be/71RML96XxCI.
The problems of getting sight back, but not necessarily perception, are
described by Michael May at http://youtu.be/bnefTJx2sCo.

For Chaps. 11 and 4

Hebb postulate illustrated at retinotectal synapses by Mu-ming Poo http://
youtu.be/hLs3m6nIJ1E.

For Chap. 12
The role of immune system molecules in visual plasticity by Carla Shatz
http://youtu.be/WFA9xbhI3yc.