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Progress in Brain Research
Volume 273
Circadian and Visual

Neuroscience
Serial Editor
Vincent Walsh
Institute of Cognitive Neuroscience
University College London
17 Queen Square
London WC1N 3AR UK
Editorial Board
Tanya Calvey, Johannesburg, South Africa

University of the Witwatersrand
Hamed Ekhtiari, USA
University of Minnesota
Chi Ieong Lau, Taipei, Taiwan
Shin Kong Wu Ho-Su Memorial Hospital
Shane O’Mara, Dublin, Ireland
Trinity College
Flavia Santos, Ireland
University College Dublin
Progress in Brain Research
Volume 273
Circadian and Visual

Neuroscience
Edited by
Nayantara Santhi
Department of Psychology,
Northumbria University,
Newcastle Upon Tyne, United Kingdom
Manuel Spitschan
Translational Sensory & Circadian Neuroscience, Max Planck
Institute for Biological Cybernetics, Tübingen, Germany
Chronobiology & Health, TUM Department of Sport
and Health Sciences (TUM SG),
Technical University of Munich, Munich;
TUM Institute for Advanced Study (TUM-IAS),
Technical University of Munich, Garching, Germany
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Contributors
Annette E. Allen
Centre for Biological Timing, Division of Neuroscience and Experimental Biology,
Faculty of Biology, Medicine and Health, University of Manchester, Manchester,
United Kingdom
Beatriz Baño-Otálora
Centre for Biological Timing, Division of Neuroscience and Experimental Biology,
Faculty of Biology, Medicine and Health, University of Manchester, Manchester,
United Kingdom
John L. Barbur
Centre for Applied Vision Research, The Henry Wellcome Laboratories for Vision
Science, School of Health Sciences, City, University of London, London,
United Kingdom
Pablo A. Barrionuevo
Instituto de Investigación en Luz, Ambiente y Visión, Consejo Nacional de
Investigaciones Cientı́ficas y T
ecnicas—Universidad Nacional de Tucumán,
Tucumán, Argentina
Peter Blattner
Federal Institute of Metrology METAS, Bern-Wabern, Switzerland; President of
the International Commission on Illumination (CIE), Vienna, Austria
David H. Brainard
Department of Psychology, University of Pennsylvania, Philadelphia, PA,
United States
Timothy M. Brown
Centre for Biological Timing, Faculty of Biology Medicine and Health, University of
Manchester, Manchester, United Kingdom
Hugo Calligaro
Salk Institute for Biological Studies, La Lolla, CA, United States
Ashley E. Copenhaver
Department of Biological Sciences, University of Maryland, Baltimore County
(UMBC), Baltimore, MD, United States
Nicolas P. Cottaris
Department of Psychology, University of Pennsylvania, Philadelphia, PA,
United States
Ouria Dkhissi-Benyahya
Univ Lyon, Universit
e Claude Bernard Lyon 1, Inserm, Stem Cell and Brain
Research Institute, Bron, France
v
vi Contributors
Greg J. Elder
Northumbria Sleep Research, Department of Psychology, Faculty of Health and
Life Sciences, Northumbria University, Newcastle upon Tyne, United Kingdom
Rhea T. Eskew, Jr.
Department of Psychology, Northeastern University, Boston, MA, United States
Dorothee Fischer
Department of Sleep and Human Factors Research, Institute for Aerospace
Medicine, German Aerospace Center, Cologne, Germany
Elisabeth Flo-Groeneboom
Department of Clinical Psychology, Faculty of Psychology, University of Bergen,
Bergen, Norway
Russell G. Foster
Sir Jules Thorn Sleep and Circadian Neuroscience Institute (SCNi), Kavli Institute
for NanoScience Discovery, Nuffield Department of Clinical Neurosciences,
University of Oxford, Oxford, United Kingdom
Jingyi He
Cassie J. Hilditch
Fatigue Countermeasures Laboratory, Department of Psychology, San Jose State
University, San Jos
e, CA, United States
Anya Hurlbert
Centre for Transformative Neuroscience and Institute of Biosciences, Newcastle
University, Newcastle upon Tyne, United Kingdom
Luis A. Issolio
ecnicas—Universidad Nacional de Tucumán;
Departamento de Luminotecnia, Luz y Visión, Facultad de Ciencias Exactas y
Tecnologı́a, Universidad Nacional de Tucumán, Tucumán, Argentina
Antonin Jandot
Univ Lyon, Universit
e Claude Bernard Lyon 1, Inserm, Stem Cell and Brain
Research Institute, Bron, France
Tara A. LeGates
(UMBC); Department of Physiology, University of Maryland School of Medicine,
Baltimore, MD, United States
Renske Lok
Department of Psychiatry and Behavioral Sciences, Stanford University, Stanford,
CA, United States
Contributors vii
Takuma Morimoto
Department of Experimental Psychology, University of Oxford, Oxford,
United Kingdom; Department of General Psychology, Justus-Liebig-Universit€at
Gießen, Gießen, Germany
Joshua W. Mouland
Centre for Biological Timing, Faculty of Biology Medicine and Health, University of
Manchester, Manchester, United Kingdom
Ruben Pastilha
Centre for Transformative Neuroscience and Institute of Biosciences, Newcastle
University, Newcastle upon Tyne, United Kingdom
Stuart N. Peirson
Carina A. Pothecary
Oscar U. Preciado
Departamento de Luminotecnia, Luz y Visión, Facultad de Ciencias Exactas y
Tecnologı́a, Universidad Nacional de Tucumán, Tucumán, Argentina
Luke L.A. Price
Centre for Radiation, Chemical and Environmental Hazards, Public Health
England, Chilton, Harwell Campus, Didcot, Oxfordshire, United Kingdom;
Secretary of Division 6 for Photobiology and Photochemistry of CIE, Vienna,
Austria
Roshae C. Roberts
(UMBC), Baltimore, MD, United States
Marı́a L. Sandoval Salinas
Instituto de Investigaciones en Biodiversidad Argentina, Facultad de Ciencias
Naturales e Instituto Miguel Lillo, Universidad Nacional de Tucumán, Tucumán,
Argentina
Ma’ayan Semo
viii Contributors
Keizo Shinomori
School of Information/Research Institute, Kochi University of Technology, Kochi,
Japan
Melissa A. St. Hilaire
Division of Sleep and Circadian Disorders, Brigham and Women’s Hospital;
Division of Sleep Medicine, Harvard Medical School, Boston, MA, United States
Laura C.E. Steel
S.K.E. Tam
Yesenia Taveras-Cruz
Selma Tir
Vladyslav V. Vyazovskiy
for NanoScience Discovery, Department of Physiology, Anatomy and Genetics,
Brian A. Wandell
Psychology Department and the Stanford Center for Image Systems Engineering,
Stanford University, Stanford, CA, United States
John S. Werner
Department of Ophthalmology & Vision Science, University of California Davis,
Sacramento, CA, United States
Jamie M. Zeitzer
Department of Psychiatry and Behavioral Sciences, Stanford University, Stanford;
Mental Illness Research Education and Clinical Center, VA Palo Alto Health Care
System, Palo Alto, CA, United States
Contents
Contributors .............................................................................................................. v
Foreword ............................................................................................................... xvii
Introduction ........................................................................................................... xxi
CHAPTER 1 Circadian and visual photometry.................................1

Luke L.A. Price and Peter Blattner
1. The basis of physical photometry .................................................. 2
1.1 Additional notes to Eqs. (1) to (7) .......................................... 4
2. Colorimetry .................................................................................... 4
3. IIL responses .................................................................................. 7
3.1 Additional notes to Eqs. (14) and (15) ................................... 9
4. Conclusions .................................................................................... 9
References......................................................................................... 10
CHAPTER 2 Optical stimulation systems for studying human

vision..........................................................................13
Pablo A. Barrionuevo, Oscar U. Preciado,
Marı́a L. Sandoval Salinas, and Luis A. Issolio
1. Introduction.................................................................................. 14
1.1 The silent substitution method .............................................. 14
2. Optical stimulation systems......................................................... 15
2.1 Nonimage-forming arrangements ......................................... 15
2.2 Image-forming arrangements ................................................ 19
3. Characterization and calibration principles ................................. 25
3.1 Stimulator response curve ..................................................... 26
3.2 Spectral power distribution................................................... 28
3.3 Additivity .............................................................................. 30
3.4 Modulation transfer function ................................................ 30
3.5 Spatial uniformity ................................................................. 30
3.6 Temporal stability ................................................................. 31
3.7 Sources of artifacts................................................................ 31
4. Conclusions .................................................................................. 32
References......................................................................................... 32
CHAPTER 3 Hyperspectral characterization of natural

lighting environments.................................................37
Takuma Morimoto
1. Introduction.................................................................................. 37
2. Method ......................................................................................... 39
ix
x Contents
2.1 Capturing images of a mirror sphere .................................... 40

2.2 Image processing ................................................................... 41
3. Application ................................................................................... 43
4. Conclusion .................................................................................... 46
Acknowledgment ............................................................................... 46
References......................................................................................... 46
Further reading .................................................................................. 48
CHAPTER 4 Endogenous functioning and light response of the

retinal clock in vertebrates.......................................49
Antonin Jandot, Hugo Calligaro, and
Ouria Dkhissi-Benyahya
1. Introduction .................................................................................. 50
2. Molecular organization of the retinal clock in vertebrates ......... 51
3. Cellular organization of the retinal clock in vertebrates ............. 53
4. Retinal circadian rhythms ............................................................ 56
5. The light response of the retinal clock........................................ 57
6. Future directions .......................................................................... 59
Acknowledgments ............................................................................. 60
References......................................................................................... 60
CHAPTER 5 Light-dependent effects on mood: Mechanistic

insights from animal models......................................71
Ashley E. Copenhaver, Roshae C. Roberts, and
Tara A. LeGates
1. Introduction .................................................................................. 72
2. Light detection pathways and effects on physiology in
mammals...................................................................................... 73
3. Effects of light on mood in humans ............................................ 74
3.1 Light at night ......................................................................... 74
3.2 Transmeridian travel ............................................................. 75
3.3 Seasonal changes in daylength ............................................. 75
3.4 Bright light therapy ............................................................... 76
4. Animal models ............................................................................. 77
4.1 Using animals to study mood disorders ............................... 78
4.2 Nocturnal versus diurnal ....................................................... 79
5. Mood alterations induced by light-dependent disruption
of circadian rhythms .................................................................... 80
6. Direct effects of light ................................................................... 84
7. Bright light therapy ...................................................................... 87
8. Discussion .................................................................................... 88
References......................................................................................... 89
Contents xi
CHAPTER 6 Rodent models in translational circadian

photobiology...............................................................97
Selma Tir, Laura C.E. Steel, S.K.E. Tam, Ma’ayan Semo,
Carina A. Pothecary, Vladyslav V. Vyazovskiy,
Russell G. Foster, and Stuart N. Peirson
1. Introduction.................................................................................. 98
2. The role of rodent models in circadian photobiology ................. 99
3. Translational circadian photobiology ........................................ 100
4. Strengths and limitations of rodent models ............................... 101
4.1 Strengths .............................................................................. 102
4.2 Limitations .......................................................................... 103
5. Future directions........................................................................ 105
6. Conclusions ................................................................................ 108
Acknowledgments........................................................................... 108
References....................................................................................... 109
CHAPTER 7 Slow vision: Measuring melanopsin-mediated

light effects in animal models.................................117
Annette E. Allen and Beatriz Baño-Otálora
1. Introduction................................................................................ 117
2. Methods of studying melanopsin at a systems level ................. 122
2.1 Eliminating rod/cone responses .......................................... 122
2.2 Manipulating melanopsin activity...................................... 123
2.3 Modulating ipRGC activity ................................................ 124
2.4 Stimulus spectra and receptor silent substitution ............... 124
3. Melanopsin contribution to nonimage-forming function:
Methods and findings ................................................................. 126
3.1 Circadian photoentrainment ................................................ 126
3.2 Pupillary light reflex ........................................................... 130
3.3 Sleep and behavioral regulation ......................................... 131
3.4 Mood regulation .................................................................. 132
4. Closing remarks......................................................................... 133
References....................................................................................... 135
CHAPTER 8 Beyond irradiance: Visual signals influencing

mammalian circadian function................................145
Joshua W. Mouland and Timothy M. Brown
1. Introduction................................................................................ 145
2. Experimental paradigms providing insight into the
photoentrainment mechanism .................................................... 146
xii Contents
3. Evidence for inner and outer retinal sources of circadian

control........................................................................................ 149
4. Retinal photoreceptor specializations ........................................ 151
5. Rod contributions to circadian photoentrainment ..................... 153
6. Cone contributions to circadian photoentrainment................... 155
7. Spatiotemporal properties of circadian responses ..................... 158
8. Do insights into rodent photoentrainment translate to
humans? ...................................................................................... 160
9. Conclusion .................................................................................. 161
References....................................................................................... 163
CHAPTER 9 Circadian photoreception: The impact of light

on human circadian rhythms...................................171
Jamie M. Zeitzer and Renske Lok
1. Phase response curve ................................................................. 172
2. Duration sensitivity .................................................................... 174
3. Intensity response curve ............................................................. 175
4. Spectral sensitivity ..................................................................... 176
References....................................................................................... 177
CHAPTER 10 Modeling (circadian)................................................181

Melissa A. St. Hilaire
1. Introduction ................................................................................ 181
2. Foundational aspects of modeling the circadian system ........... 182
2.1 Process C: A simple sinusoid............................................. 182
2.2 Oscillator models of the mammalian circadian system .... . 183
3. Representation of light in models of the circadian system ....... 185
3.1 Power-law relationship for circadian light effects............. 185
3.2 Process L: The light preprocessor ...................................... 186
3.3 Beyond lux as light input to mathematical models
of the circadian system ....................................................... 187
3.4 Temporal dynamics of light and cross-species
comparisons ......................................................................... 189
4. Application of mathematical models to improve human
health and performance .............................................................. 191
4.1 Predictions of neurobehavioral performance and
alertness ............................................................................... 191
4.2 Individualized predictions of circadian phase .................... 192
4.3 Generating testable hypotheses for light treatment
and future experiments........................................................ 193
5. Conclusions ................................................................................ 194
References....................................................................................... 195
Contents xiii
CHAPTER 11 Visual encoding: Principles and software..............199

Brian A. Wandell, David H. Brainard, and
Nicolas P. Cottaris
1. Introduction................................................................................ 199
2. Representing the scene radiance ................................................ 200
2.1 Planar scenes: Calibrated display ....................................... 201
2.2 Planar scenes: Natural image approximation ..................... 202
2.3 Display variation ................................................................. 204
2.4 Three-dimensional scenes................................................... 206
3. Optics: The retinal irradiance.................................................... 208
3.1 Image formation: 2D ........................................................... 209
3.2 Image formation: 3D ........................................................... 213
4. Light transduction...................................................................... 215
4.1 Rod and cone photoreceptors .............................................. 215
4.2 Intrinsically photosensitive retinal ganglion cells .............. 224
5. Example calculations ................................................................. 226
6. Summary.................................................................................... 226
References....................................................................................... 227
CHAPTER 12 Visual psychophysics: Luminance and color..........231

Yesenia Taveras-Cruz, Jingyi He, and Rhea T. Eskew, Jr.
1. Introduction................................................................................ 231
2. Psychophysical methods ............................................................ 233
2.1 Classical methods ................................................................ 235
2.2 Signal detection theory ....................................................... 237
2.3 Adaptive methods ............................................................... 238
3. Selected results ........................................................................... 239
3.1 Luminous efficiency and spectral sensitivity ..................... 239
3.2 Color vision ......................................................................... 242
3.3 Spatial and temporal sensitivity: Effects of light level ...... 247
4. Final thoughts & resources........................................................ 253
4.1 Additional resources........................................................... 253
References....................................................................................... 254
CHAPTER 13 Aging of visual mechanisms....................................257

Keizo Shinomori, John L. Barbur, and John S. Werner
1. Introduction................................................................................ 257
2. Optical changes associated with aging ...................................... 258
2.1 Senescent changes in the eye pupil .................................... 258
2.2 Senescent changes in preretinal filters ............................... 259
2.3 Senescent changes in retinal image quality........................ 261
xiv Contents
3. Senescent changes in sensitivity of rod and cone pathways ..... 262

3.1 Dark adaptation and rod sensitivity .................................... 262
3.2 Age-related changes in cone sensitivity ............................. 263
3.3 Color discrimination ........................................................... 263
3.4 Spatial vision ....................................................................... 265
3.5 Estimating sensitivity changes of ipRGCs......................... 266
3.6 Temporal vision .................................................................. 268
4. A note about unconscious visual processes ............................... 270
References....................................................................................... 271
CHAPTER 14 Seeing and sensing temporal variations in

natural daylight........................................................275
Ruben Pastilha and Anya Hurlbert
1. Measurements of natural illumination over time ...................... 277
1.1 The chromaticities of daylight ............................................ 277
1.2 Temporal variations in daylight chromaticity and
illuminance from dawn to dusk .......................................... 280
2. The dual human response to light ............................................. 287
2.1 Responses to light in nonvisual pathways.......................... 287
2.2 The role of S cone pathways .............................................. 290
3. Measuring perception of temporal variations in illumination ... 292
3.1 Visual discrimination of temporal changes in
illumination chromaticity .................................................... 292
3.2 Perceptual thresholds compared with natural
illumination temporal change characteristics..................... 295
3.3 Nonvisual and visual mechanisms underlying the
discrimination of temporal changes in illumination ........... 296
3.4 Implications and open questions......................................... 297
Acknowledgments ........................................................................... 297
References....................................................................................... 298
CHAPTER 15 Light in ecological settings: Entrainment,

circadian disruption, and interventions..................303
Dorothee Fischer and Cassie J. Hilditch
1. Introduction ................................................................................ 303
2. Entrainment in humans: Light as the dominant zeitgeber ........ 305
2.1 Models of entrainment ........................................................ 305
2.2 Chronotypes ........................................................................ 308
3. Circadian disruption: How misaligned light-dark cycles
can disrupt the circadian system ................................................ 309
3.1 Sleep regularity................................................................... 311
Contents xv
3.2 Position within a time zone ................................................ 313

3.3 Daylight saving time ........................................................... 313
3.4 Shift work ............................................................................ 314
4. Field studies: Using light to mitigate effects of circadian
disruption in ecological settings ................................................ 314
4.1 Phase shifting effects of light ............................................. 315
4.2 Zeitgeber strengthening and acute alerting properties
of light ................................................................................. 317
4.3 Non-24-h light-dark cycles ................................................. 318
4.4 Challenges and recommendations ...................................... 319
5. Conclusion .................................................................................. 322
References....................................................................................... 322
CHAPTER 16 How can light be used to optimize sleep and
health in older adults?.............................................331
Greg J. Elder and Elisabeth Flo-Groeneboom
1. Introduction................................................................................ 331
2. Normal age-related changes to sleep and circadian
rhythmicity ................................................................................. 333
2.1 Sleep.................................................................................... 333
2.2 Circadian rhythmicity ......................................................... 336
3. Can light improve sleep in older adults? ................................... 337
3.1 Bright light treatment .......................................................... 338
3.2 How might light improve circadian rhythmicity? .............. 339
3.3 How might light improve sleep? ........................................ 339
4. Methodological considerations for the use of light in sleep
and health ................................................................................... 340
4.1 What is the most effective mode of delivery? ................... 340
4.2 What is the most feasible mode of delivery? ..................... 341
4.3 What is the optimal dose? .................................................. 342
5. Can light be used to treat clinical problems? ............................ 344
5.1 Physical health .................................................................... 344
5.2 Sleep problems .................................................................... 344
5.3 Mood and depression.......................................................... 346
5.4 Circadian rhythm disorders ................................................. 346
5.5 Prevention ........................................................................... 347
6. Conclusion .................................................................................. 347
References....................................................................................... 348
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Foreword
One who kindles the light of awareness within gets true light.
—Rig Veda, circa 1500–1000 BCE
And God said, Let there be light: and there was light.
—Genesis, circa 550 BCE
All the fifty years of conscious brooding have brought me no closer to answer the
question, ’What are light quanta?’ Of course, today every rascal thinks he knows
the answer, but he is deluding himself.
—Albert Einstein, 1951 CE
Light is miraculous. Light sustains our food chain, illuminates our world, and quietly
controls the chemical tides in our bodies. Humans are predominantly a visual spe-
cies; our moment-to-moment waking consciousness is filled with the sense of vision.
It has been estimated that more than half of the central nervous system is involved in
the primary and secondary processing of vision and visual reflexes. It is no wonder,
then, that exploring the nature of light and color has been a passion for prophets, mys-
tics, philosophers, and scientists across the millennia. Brilliant minds, from Plato to
Goethe and from Newton to Einstein, have puzzled over the mystery of light and its
interaction with the human eye (Zajonc, 1993).
The central thread of this book is the interaction of light and the eye. It synthe-
sizes light both as a stimulus for the sensory capacity of vision and for the regulation
of circadian, neuroendocrine, and neurobehavioral physiology. Dr. Spitschan and
Dr. Santhi are to be congratulated for fostering the diverse voices responsible for
the collected chapters comprising this volume. Importantly, they opened the door
to many investigators in the earlier stages of their careers for contributing most of
the manuscripts. The future of any field of empirical research always depends
on an emerging cadre of investigators who bring fresh questions, innovative
approaches, and novel insights. To bring balance and diversity of perspectives,
Dr. Santhi and Dr. Spitschan have judiciously included contributions from some
more senior investigators. Astonishingly, this field of light research is founded on
frogs and swimming ghosts.
More than a century ago, studies on the Northern leopard frog, Rana pipiens,
ultimately led to the isolation and chemical characterization of the pineal hormone,
melatonin. In 1915, Carey McCord, MD, and his assistant Floyd Allen conducted
groundbreaking experiments feeding R. pipiens tadpoles with various bovine tissues.
To their surprise, tadpoles fed with a mixture of fresh plant food and desiccated
pineal tissue lost their dark pigmentation. In an interview, McCord recounted how
xvii
xviii Foreword
Allen excitedly reported: “They looked like ghosts swimming around in the water
with very black eyes!” Working from this stunning observation, McCord and Allen
conducted controlled studies on 12,000 tadpoles, thereby establishing the influence
of the pineal gland tissue on frog pigmentation (McCord and Allen, 1917). Progress
halted abruptly when McCord joined the National Guard to serve in a coordinated
mission between the United States and Mexico to stop the notorious rebel Pancho
Villa. After completing his military service, McCord returned to have a prominent
career in industrial medicine.
After retirement, McCord had a chance encounter with dermatologist Aaron
Lerner, MD, PhD, at the University of Michigan in 1952. They happened to sit to-
gether for lunch in the school cafeteria. McCord captivated Lerner’s attention when
their discussion turned to McCord’s earlier frog skin studies. The two scientists met
several times to discuss the possibility of isolating the active skin-lightening agent
from the pineal gland. Lerner later moved to Yale University where he and his
colleagues embarked on that very project.
Over a 4-year period, Lerner’s team fractioned and tested more than 250,000 cow
pineal glands in an in vitro assay system of R. pipiens frog skin cells. After 3 years,
they hit a significant roadblock. The team was in upheaval as they realized they
would need a million more glands to isolate a pure hormone from the pineal gland.
Discouraged, they gave the project one more month to provide a good lead before
ceasing the work entirely. One week before shutting down the project, Lerner had
an intuitive flash about the exact structure of the pineal hormone. His insight led
them to isolate a miniscule batch of melatonin—an invisible layer in the bottom
of an otherwise empty flask. In their frog skin assay, however, it was 100,000 times
more potent than other known skin-lightening agents such as noradrenaline and ace-
tylcholine. This crowning achievement brought the arduous, 4-year project to a suc-
cessful conclusion (Lerner et al., 1958).
Decades later, the discovery of melanopsin was also predicated on results from
studies on the frog skin. Mark Rollag, PhD, and his colleagues chose to work with a
bioassay developed from the frog Xenopus laevis, commonly named the African
clawed frog. Rollag’s team was interested in identifying the melatonin receptor
and melatonin’s capacity to regulate cell physiology (White et al., 1987). Those stud-
ies employed an in vitro Xenopus melanophore assay. A chance observation noted
that this assay responded to very subtle differences of light and shadow on their
lab bench, thus highlighting its utility in the study of extra-retinal photoreception.
In 1996, Ignacio Provencio, PhD, a scientist with a background in circadian photo-
reception, joined Drs. Rollag and Guisen Jiang as a postdoctoral fellow. They set out
to identify the active photopigment in frog skin melanophores. This quest was
facilitated by employing a Xenopus melanophore cDNA library screen, which
proved invaluable for narrowing the search for the active photopigment in tadpole
skin cells. After 2 years of intensive work, the project successfully culminated in
the identification and molecular characterization of the photopigment melanopsin.
In a collaboration with William Par Hayes, PhD, the team demonstrated through
in situ hybridization that melanopsin was localized in frog skin melanophore cells
Foreword xix
as well as in the frog retina, iris, and hypothalamic nuclei. Those results suggested
that melanopsin played a role in circadian physiology as well as in the photic control
of frog skin pigmentation (Provencio et al., 1998).
Ultimately, photosensitive frog skin was pivotal in the discovery of two key
molecules, the hormone melatonin and the photopigment melanopsin. Each of these
molecules is cited numerous times across the chapters in this volume.
Dr. Santhi and Dr. Spitschan have curated a collection of contributions woven
from both fresh and familiar voices. This book is an invaluable tool for understanding
our present view of how light stimulates the retina to support vision and physiolog-
ical regulation. It is also a portal to the future of this rapidly developing field.
George C. Brainard, PhD
Professor of Neurology
Director, Light Research Program
Thomas Jefferson University
Philadelphia, Pennsylvania
References
Lerner, A.B., Case, J.D., Takahashi, Y., Lee, Y., Mori, W., 1958. Isolation of melatonin, the
pineal gland factor that lightens melanocytes. J. Am. Chem. Soc. 80, 2587.
McCord, C.P., Allen, F.B., 1917. Evidence associating pineal gland function with alterations
in pigmentation. J. Exp. Zool. 23, 207–224.
Provencio, I., Jiang, G., De Grip, W.J., Hayes, W.P., Rollag, M.D., 1998. Melanopsin: an opsin
in melanophores, brain, and eye. PNAS 95, 340–345.
White, B.H., Sekura, R.D., Rollag, M.D., 1987. Pertussis toxin blocks melatonin-induced pig-
ment aggregation in Xenopus dermal melanophores. J. Comp. Physiol. B 157, 153–159.
Zajonc, A., 1993. Catching the Light: The Entwined History of Light and Mind. Oxford
University Press, New York.
Introduction
Quite apart from vision, light exerts a potent nonvisual influence on physiological
and behavioral functions including sleep, circadian rhythms, mood, and alertness.
Until the late 1990s, rods and cones were considered to be the eye’s sole light
detectors, with the former considered to predominate nighttime vision and the latter
color vision. The early challenge to the idea that rods and cones account for all of the
eye’s light functions came from the intriguing observations that the sleep-wake
cycle of some functionally blind individuals tracked the solar day and that these
individuals also exhibited light-induced effects on the melatonin section. At the same
time, animal studies showed similar results, where mice engineered to have no rods
or cones exhibited circadian entrainment just as the visually blind humans did. These
converging lines of evidence finally led to the seminal discovery of a third photore-
ceptor system, the intrinsically photosensitive retinal ganglion cells (ipRGCs), capa-
ble of producing light-induced physiological effects. We now know that the visual
and nonvisual influences of light on physiology and behavior are mediated through
complex interactions between the rods, cones, and ipRGCs. Yet, explorations of the
visual and nonvisual effects of light proceed independently. In this volume, we bring
together researchers from these diverse areas to showcase the landscape of light
effects on our physiology and behavior.
In his foreword to this volume, George Bud Brainard eloquently lays out our
motivation and rationale for putting together this collection. Drs. Blattner and Price
discuss the current standards for both circadian and visual photometry, explaining
the definitions and rationale behind the computation involved in these standards.
Investigations of the visual and nonvisual effects of light require exacting and precise
methodologies, particularly with regard to stimulating the light receptors in the eye.
Drs. Barrionuevo, Preciado, Sandoval Salinas, and Issolio walk us through the com-
plex arrangements required to achieve optical stimulation of the visual and nonvisual
receptors and pathways. Recent explorations of light and physiology and behavior
have expanded to consider our ambient light environment and daily light exposure
patterns, beyond the laboratory milieu. In his chapter, Dr. Morimoto discusses the
intricacies of characterizing light in real-world environments.
Animal models have provided the initial insights into the neural pathways and
networks underpinning the influences of light, visual and nonvisual, on physiology
and behavior. Drs. Dkhissi-Benyahya, Jandot, and Calligaro discuss the molecular
machinery and retinal clocks that lie at the heart of circadian photoentrainment. Cir-
cadian rhythmicity is not the only nonvisual function of light. Light appears to have
independent effects on other aspects of biology including mood and alertness. The
chapter by Drs. LeGates, Copenhaver, and Roberts provides a comprehensive discus-
sion on the neuronal mechanisms underlying the direct effect of light on mood.
Drs. Peirson, Steel, and Tir explain how animal models have informed our under-
standing of light effects in humans. At the heart of the nonvisual effect of light lies
the melanopsin-driven role of ipRGCs, which is discussed in detail in the chapter by
xxi
xxii Introduction
Drs. Allen and Baño-Otalora. While the ipRGCs are vital to circadian photoentrain-
ment, they do convey signals from the rods and cones to the circadian clock, as
shown and discussed by Drs. Mouland and Brown in their chapter.
Moving onto human studies, the characteristics of the light stimulus including its
timing, duration, intensity, and spectral composition modulate the strength of the
nonvisual effect of light. Drs. Zeitzer and Lok present a comprehensive summary
of the decades of research on how these aspects of light influence circadian photo-
entrainment and other aspects of physiology and behavior. The efficacy of practical
and clinical applications of light interventions rests on our ability to accurately model
findings in the literature. Dr. St. Hilaire, in her chapter, focuses on the mathematical
models of circadian rhythms that incorporate light as a stimulus. Dr. Wandell, in his
chapter, discusses the principles and parameters of optics and visual encoding and
presents an open-source toolbox that estimates these parameters. Drs. Taveras-Cruz,
He, and Eskew Jr. present an eloquent discussion on the visual psychophysics, a
chapter complementary to the circadian photoreception chapter by Drs. Zeitzer
and Lok. Precision medicine depends on the discovery of factors contributing to
individual differences in physiology and behavior. Age is one of the strongest con-
tributors to individual variation in physiology. Drs. Werner, Barbur, and Shinomori
review our current understanding of the aging of visual mechanisms. We conclude
this volume with three chapters that elucidate the light influences and applications in
the real world. Drs. Pastilha and Hurlbert discuss how the eye senses changes in
natural illumination and how this shapes our physiology.
Drs. Fischer and Hilditch take this thinking a step further in their chapter by
showing us how our daily light exposure in the real world leads to circadian disrup-
tion and how we can use light as a countermeasure against it. As we age, our sleep
and circadian rhythms are affected. Drs. Elder and Flo-Groeneboom provide insights
into the state-of-the-art development in light interventions and strategies to optimize
sleep and circadian rhythms in older adults, both in health and in disease.
It has been a great delight to bring together such an exciting and diverse group of
researchers to present their work and understanding of how light influences our daily
lives constantly. We hope this volume will serve in bringing together the ideas and
discoveries of vision science and circadian physiology to develop novel hypotheses
and new questions. We thank all the authors for their invaluable contributions to this
volume and the publishing team for their patience and help. And last but not least, we
thank George Brainard for the very poetic foreword to this volume.
Nayantara Santhi
Manuel Spitschan
CHAPTER
Circadian and visual

photometry
1
Luke L.A. Pricea,b,∗ and Peter Blattnerc,d
a
Centre for Radiation, Chemical and Environmental Hazards, Public Health England, Chilton,
Harwell Campus, Didcot, Oxfordshire, United Kingdom
b
Secretary of Division 6 for Photobiology and Photochemistry of CIE, Vienna, Austria
c
Federal Institute of Metrology METAS, Bern-Wabern, Switzerland
d
President of the International Commission on Illumination (CIE), Vienna, Austria
∗
Corresponding author: e-mail address: luke.price@phe.gov.uk
Abstract
Photometry is the metrology of light—optical radiation seen by the human eye due to its
action on retinal photoreceptors. Its origins are closely tied to the International Commission
on Illumination (CIE), which remains responsible for photometry standards and the language
of light used in science and technology.
When in 1931 it had become possible to model the response to light of the human eye based
on reliable spectroradiometry data, the CIE published standard formulae for predicting the
luminance of a stimulus. These and related colorimetry formulae are still in use, having been
internationally agreed and adopted. Both fields continue to be the subject of active research
and increasing accuracy.
CIE S 026:2018 represents another milestone for the metrology of light (CIE, 2018a). It is
the first standard where light is considered for its ability to evoke circadian and neurophysi-
ological responses, and includes the spectral sensitivity of melanopsin—a retinal photopig-
ment discovered, and shown to be contributing to and influencing responses from human
intrinsically-photosensitive retinal ganglion cells (ipRGCs), only 20 years ago (Berson
et al., 2002; Hattar et al., 2002; Provencio et al., 1998). These accessory visual functions also
depend to some extent on inputs from the rods and three types of cones; until very recently,
rods and cones (or “classical photoreceptors”) were the only photoreceptors in visual models.
If photometry standards are replaced with modern physiological data, consistent changes
should be expected in the photometry of these accessory functions.
This chapter outlines the current standards, their definitions and calculations, and how the
main elements are related.
Progress in Brain Research, Volume 273, ISSN 0079-6123, https://doi.org/10.1016/bs.pbr.2022.02.014

1
2 CHAPTER 1 Circadian and visual photometry
Keywords
Photometry, Colorimetry, Circadian, Melanopsin, Non-visual, Photobiology, Spectroradio-
metry, Standards
1 The basis of physical photometry

In order to quantify and compare photobiological and photochemical effects, appro-
priate metrics (i.e., measured quantities and physical units) have to be agreed. Since
1960 the International System of Units, the SI, has been used around the world as the
preferred system of units and the basic language for science, technology, industry
and trade. It was established by a resolution at the 11th meeting of the Conference
Generale des Poids et Mesures, the CGPM (known in English as the General
Conference on Weights and Measures).
The present SI continues to reflect the historical importance of human vision in
measuring light, and the reproducible physical quantities that replace the subjective
visual image-forming sensitivity of the human eye are expressed using specialized
photometric units, namely the candela, lumen and lux, having the symbols cd, lm and
lx, respectively. The link between the photometric units and the other physical units
is made by the defining constant Kcd (BIPM, 2019a): according to the definition, the
luminous efficacy of monochromatic radiation of frequency 540 THz is exactly
683 lm/W.
In practice, however, humans and their surroundings are very rarely illuminated
by monochromatic radiation of 540 THz (which corresponds to about 555 nm and is
perceived as “green” in colour). Appendix 2 of the SI brochure (BIPM, 2019b) there-
fore defines how the photometric radiation equivalence of polychromatic radiation is
calculated. Depending on the observer’s viewing conditions, different spectral lumi-
nous efficiency functions Vx(λ) are defined. These functions are usually normalized
to unity at the wavelength of highest sensitivity. As set out in Eqs. (1) and (2), the
luminous flux for a given observer Φv,x is given by a spectrally-weighted summation
of the optical radiation Φe,x(λ), multiplied by Kcd, and divided through by the value of
the corresponding spectral luminous efficiency function Vx(λcd) at the wavelength in
standard air, λcd corresponding to the frequency of 540 THz in standard air:
ð
Φv,x ¼ Km Φe,λ ðλÞ Vx ðλÞ dλ (1)
λ
K m ¼ K cd =V ðλcd Þ ¼ 683:002 lm=W (2)
K 0 m ¼ Kcd =V 0 ðλcd Þ ¼ 1,700.05 lm=W (3)
The spectral luminous efficiency function most relevant in practice was pub-
lished in 1931 by the International Commission on Illumination CIE (CIE, 2019a)
and applies to a “typical” observer adapted to daylight (photopic vision, Eq. (2)),
with a narrow field of view (about 4° or less). In addition, CIE has published several
other standard observers, including for scotopic (Eq. (3), with K’m substituting for
1 The basis of physical photometry 3
1750 1700.048 lm/W Photopic

Scotopic
Luminous eﬃcacy, K / (lm/W) 1500
Maxima
1250
1000
750 683.002 lm/W
500
250
0
400 450 500 550 600 650 700
wavelength, λ / nm
FIG. 1
The CIE photopic and scotopic spectral luminous efficacies of radiation, K(λ) and
K0 (λ)—dashed black and light gray lines—and their respective maxima as diamonds.
Luminous efficacies agree exactly at the frequency of 540 1012 Hz (SI definition),
corresponding to λcd, approximately 555.017 nm in standard air (close to the photopic
maximum, λmax 555 nm). Rhodopsin is solely responsible for the scotopic spectral luminous
efficacy, so that srh(λ) ¼ V 0 (λ) ¼ K0 (λ)/K0 (λmax), where λmax 507 nm.
Km in Eq. (1)), mesopic vision (see CIE, 2010) and for situations where the visual
target has an angular subtense larger than 4° or is seen off-axis (CIE, 2005). These
“observers” relate to different visual conditions and visual fields (e.g., see the
additional notes below), and all represent a person with typical visual responses
at approximately 32 years of age. Fig. 1 shows the dependence on wavelength of
the luminous efficacy functions for vision in photopic and scotopic conditions,
K(λ) and K0 (λ), i.e., Km V(λ) and K0 m V0 (λ), respectively.
Photobiological or photochemical responses to optical radiation that are suffi-
ciently distinct from the human image-forming visual sensation are not described
by standard photometry and are instead dealt with using the radiometric system
codified in Appendix III of the SI Brochure (BIPM, 2020). Several photobio-
logical responses have their in vivo spectral sensitivity standardized as an action
spectrum, allowing calculation of weighted radiometric quantities, according to
Eqs. (4) and (5). ð
Φx ¼ Φe,λ ðλÞ sx ðλÞ dλ (4)
λ
ð
Φp,x ¼ Φp,λ ðλÞ sp,x ðλÞ dλ (5)
λ
These equations, (4) and (5), are valid for the two main branches of SI radiom-
etry known as the energy system (see Section 1.1) and the photon system,
respectively. The existence of two radiometric systems has in the past led to con-
fusion about the difference between the action spectra in each system for the same
photobiological response. This has been resolved in Appendix III that was
released with the 9th Edition of the SI Brochure (see BIPM, 2020). Eq. (6) governs
the relationship between the energy and photon systems’ action spectra for a
response x. Eq. (6) then sets out how to determine the renormalization
constant, γ x. However, a much easier way to renormalize when using a computer
to convert an action spectrum from one system to another, is to divide the con-
verted function throughout by its maximum value (after using any convenient
value for γ x), in terms of the frequency of light, f, and using two of the fundamental
constants of nature h and c also defined in the SI Brochure.
sp,x ðλÞ ¼ γ x sx ðλÞ h f ¼ γ x sx ðλÞ h c=ðnðλÞ λÞ (6)
1=γ x ¼ max ½ sx ðλÞ h f (7)
1.1 Additional notes to Eqs. (1) to (7)

In Eq. (1) the unit symbol for the luminous flux, Φv,x, is lm. In general, the units of
photometry are common to all the visual observer conditions, and there is no such
thing as a photopic, scotopic or mesopic lumen. In Eqs. (2) and (3), it is common
practice to use the approximations Km Kcd ¼ 683 lm/W and K0 m 1,700 lm/W.
For historical reasons, the energy system in Eq. (4) is almost universally referred
to simply as the radiometric system (including in the SI Brochure’s Appendix III),
and the use of subscript “e” to denote that quantities and action spectra belong to this
system is only required for spectral quantities, or for additional clarity. The term
“energy system” is used here to avoid the ambiguity this often creates, especially
when comparting the two systems. The unit symbols for the “energy” and photon
radiant fluxes in Eqs. (4) and (5) are W and s1, respectively; the number of photons
is dimensionless. As with photometric units, these same units apply for all different
responses (other than photometric responses).
Eqs. (6) and (7) define a subtle difference between the realization of the action
spectrum in the two systems for a given response. The difference is greater the
broader the peak of the action spectrum is; the difference in maximum wavelength
is approximately 1 to 4 nm for opsin-based responses. The relationship between fre-
quency, f, and wavelength, λ, depends on the refractive index of the medium, n. For
simplicity, the refractive index of standard air is usually assumed to be constant
across the visible spectrum, with na(λcd) 1.00028, so that hf hc/1.00028λ.
2 Colorimetry
A concept similar to photometry can be applied for quantifying colour perception:
in colorimetry, colour-matching functions are used to calculate triplets of numbers
(also called tristimulus values) from a measured colour stimulus (CIE, 2018b).
2 Colorimetry 5
The most commonly used system is the CIE XYZ trichromatic system, called the
CIE 1931 standard colorimetric system (CIE, 2006, 2019b). This is based on
the CIE 1931 colour-matching functions, that define the colour-matching properties
of an average observer with normal colour vision when viewing fields of angular
subtense between 1° and 4°, denoted by x(λ), y(λ) and z(λ) the corresponding tris-
timulus values X, Y and Z.
Eq. (8) for X, has corresponding versions for Y and Z that use the same constant k,
the same spectral quantity, here Φe,λ(λ), and substitute functions y (λ) and z (λ),
respectively for x(λ): ð
X ¼ k Φe,λ ðλÞ xðλÞ dλ (8)
λ
By construction of the CIE 1931 system y(λ) equals the V(λ) function, so a typical
choice is to use k ¼ Km and a spectral irradiance Ee,λ(λ) as the spectral quantity,
yielding a value of Y equal to illuminance.
The chromaticity coordinates (x, y) define the chromaticity of a visual stimulus,
which is given by its relative spectral distribution. The coordinates are normalized to
the sum of the tristimulus values:
x ¼ X=ð X + Y + Z Þ (9)
y ¼ Y=ð X + Y + Z Þ (10)
Having specified (x, y), z becomes redundant:

z¼1xy (11)
For larger field of views, the CIE 1964 standard colorimetric observer is defined
in (CIE, 2019b), complementary to the 1931 system.
Since colorimetry was established in 1931, there have been improvements in the
metrology of colour stimuli and important advances in the knowledge of colour
vision. The sensation of colour results from physiological processes, starting with
the capture of photons by the cones in the retina. The fundamental sensitivities of
the cones in vivo need to be precisely known to accurately specify a colour stimulus
from a given spectral power distribution.
In 2006 CIE published the relative spectral sensitivities of cone fundamentals for
the long-wave sensitive (LWS), middle-wave sensitive (MWS) and short-wave sen-
sitive (SWS) cones for a typical observer as measured at the entrance of the eye (CIE,
2006). They were derived from the most comprehensive experimental color-
matching data collected to date and allow a parameterization in respect to both
the field of view and age of the observer. Fig. 2 shows these action spectra in the
radiometric system.
A cone-fundamental as redundant spectral luminous efficiency function
VF(λ) can be defined as a linear combination of the cone fundamentals l(λ), m(λ)
and s(λ) as follows:
VF ðλÞ ¼ α lðλÞ + β mðλÞ + γ sðλÞ (12)
1 L-cone
M-cone
0.8
Relative efficiency
S-cone
0.6
0.4
0.2
0
400 450 500 550 600 650 700
wavelength, λ / nm
FIG. 2
The 2° cone fundamentals for the long-wave sensitive (LWS, light gray line), middle-wave
sensitive (MWS, mid gray line) and short-wave sensitive (SWS, black line) cones, denoted
l (λ), m(λ) and s(λ), respectively.
where α, β and γ are weighting constants. It is however a basic assumption that the
short-wave-cone system does not notably contribute to luminance, thus reducing
Eq. (12) to a linear combination of l(λ) and m(λ). The determination of the weighting
constants is outlined in (CIE, 2015a), the respective values are included in the
reduced equation as follows:
VF ðλÞ ¼ 0:689 902 72 lðλÞ + 0:348 321 89 mðλÞ (13)
VF(λ) is normalized to a peak at 556.1 nm and as redundant is shown in Fig. 3.
An equivalent cone-fundamental system has been defined for a 10° field size, with
corresponding action spectra given by defining corresponding action spectra l10(λ),
m10(λ) and s10(λ) (see CIE, 2015a).
To adopt the CIE cone-fundamental systems as an international standard would
have wide reaching implications for science and industry for both colorimetry and
photometry, and CIE Technical Committee 1-98 was set up to define a
“roadmap” for developing a new self-consistent system of CIE colorimetry. The suc-
cess of this project is likely to depend on whether the improvements are considered
sufficiently important by various stakeholders to justify the costs involved in switch-
ing to a new system of colorimetry and photometry.
Whereas the aforementioned 1964 functions were obtained by asking subjects to
disregard the central 2° field, the current functions allow for the annular sensitivity to
be established at 1° intervals up to 10°.
To ensure consistency within CIE documents governing photometry, colorimetry
and IIL responses (see Section 4 for definition), the action spectra introduced in the
3 IIL responses 7
1
Relative luminous efficiency
0.1
0.01
10° cone fundamentals

0.001
2° cone fundamentals
CIE photopic standard
0.0001
400 450 500 550 600 650 700
wavelength, λ / nm
FIG. 3
The CIE photopic luminous efficiency of radiation shown as a semi-log plot: V(λ) (“CIE
photopic standard,” dashed line) and two proposed alternatives VF(λ) (“2° cone
fundamentals,” solid dark line) and VF,10(λ) (“10° cone fundamentals,” pale solid line) based
on the cone fundamentals for a 2° and 10° field, respectively (Stockman et al., 1999;
Stockman and Sharpe, 2000).
next section as slc(λ), smc(λ) and ssc(λ) were defined to be identical to l10(λ), m10(λ)
and s10(λ), respectively, and srh(λ) is identical to V0 (λ), which is the scotopic spectral
luminous efficacy described earlier.
3 IIL responses
IIL responses stands for “ipRGC-influenced responses to light,” and are also known as
non-visual or non-image-forming responses to light (CIE, 2018a). Light arriving at the
retina, having passed through the lens and other ocular media, not only stimulates
luminous and colour vision, but influences several circadian and neurophysiological
functions (Berson et al., 2002; Hattar et al., 2002; Provencio et al., 1998; Lucas et al.,
2014). For instance, these so-called non-visual effects include the circadian rhythm
phase-shifting, suppression of pineal nocturnal melatonin secretion and the pupil
reflex. Melanopic phototransduction in the intrinsically-photosensitive retinal gan-
glion cells (ipRGCs) plays a role in addition to the signals from the cones and rods,
often appearing to take the dominant role (e.g., Brown, 2020).
Photobiological action spectra have been published in the 2018 international
standard “CIE S 026” for the IIL responses to optical radiation in the visible spectrum
that is seen as light (CIE, 2018a). The standard allows for the most agnostic inter-
pretation possible, that any one of the five photoreceptor types—short, medium
S-cone-opic
1
M-cone-opic
0.8 L-cone-opic
Relative efficiency
Rhodopic
0.6 Melanopic
0.4
0.2
0
400 450 500 550 600 650 700
wavelength, λ / nm
FIG. 4
The five CIE alpha-opic (α-opic) action spectra, sα(λ): the S-cone-opic action spectrum,
ssc(λ)—black line; the M-cone-opic action spectrum, smc(λ)—dark gray; the L-cone-opic
action spectrum, slc(λ)—light gray; the rhodopic action spectrum, srh(λ)—dashed dark gray;
and the melanopic action spectrum, smel(λ)—dashed light gray.
and long wavelength cones, rods and ipRGCs—can contribute to IIL responses, and
they have five different spectral sensitivities, sα(λ), shown in Fig. 4.
The SI versions of Eqs. (6) and (7) follow the example of the irradiance toolboxes
designed for conversions between either radiometry system and a new system of
“equivalence photometry” specifically designed for IIL responses (CIE, 2015b;
Lucas et al., 2014). These early toolboxes were superseded by the online CIE Tool-
box implementing CIE S 026 (CIE, 2018c).
The CIE S 026 Toolbox is primarily designed to support calculations of optical
radiation quantities relating to IIL responses, using calculations and convers-
ions according to Eqs. (14) and (15) for the five α-opic equivalent daylight
(D65) illuminances, ED65
v,α , or “α-opic EDIs” in lx, and the five corresponding efficacy
constants called the α-opic efficacies of luminous radiation (ELR) for daylight D65,
KD65
α,v , or “daylight α-opic ELRs”:
ð
ED65
v,α ¼ Ee,λ ðλÞ sα ðλÞ dλ=K D65
α,v (14)
λ

sc,v , K mc,v , K lc,v , K rh,v , K mel,v ¼
K D65 D65 D65 D65 D65
(15)
ð0:8173, 1:4558, 1:6289, 1:4497, 1:3262Þ mW=lm
It is important to understand that the metrology in CIE S 026 does not give in-
formation about how to predict end-point physiological responses to light, but deals
only with quantifying light for its ability to stimulate the five photoreceptor-types
and their individual sensitivities to light (i.e., for the short, medium and long
4 Conclusions 9
wavelength cones, rods and ipRGCs). The first Manchester Workshop in 2013 led to
the review paper which recommended this type of metrology (Lucas et al., 2014);
however, the second Manchester Workshop in 2019 has recently recommended that
daytime light levels—expressed as melanopic EDI—should satisfy ED65 v,mel > 250 lx,
evening light levels should satisfy ED65
v,mel < 10 lx and at night light levels should sat-
isfy ED65
v,mel < 1 lx, based largely on interpreting the results from a meta-review of
physiological data (Brown et al., 2022).
As well as using melanopic EDI, and thereby following the intrinsic response of
ipRGCs, the measurements for IIL responses and the Brown recommendations
should be made in the vertical plane of the eyes (at the appropriate position, imme-
diately in front of the eye or eyes). For IIL responses in general, the measurement
direction should concur with the direction of view, and ideally should include only
the field of vision, i.e. only light that would enter the pupil(s). This contrasts with
most measurements in visual photometry, where the stimulus is often specified in
terms of the light incident on an object with a known position and spectral properties
(e.g., reflectance). For instance, many traditional interior lighting standards refer to
the photopic illuminance in the horizontal plane of a work surface. Further details of
what researchers and lighting professionals should disclose and/or specify to deter-
mine the potential IIL responses of a given measurement can be found in the CIE and
academic literature (CIE, 2015b, 2018a; Lucas et al., 2014; Spitschan et al., 2019).
The luox Toolbox is a recent additional application designed to calculate α-opic
quantities from spectral data, which helps to generate the disclosures needed in re-
search publications (Spitschan et al., 2021).
3.1 Additional notes to Eqs. (14) and (15)

Eq. (14) defines “equivalence photometry” and is distinct from Eq. (1) for actual
photometry because the spectral distribution of a reference illuminant must be spec-
ified. When using CIE S 026:2018 (CIE, 2018a) the reference source must be the
standard daylight spectrum D65, and Eq. (15) defines the D65 efficacy constants
that, in function, are similar to Km, but are codified with inverted units. In Eq.
(14) the units for α-opic EDI, ED65
v,α , is lx, and as with photometric units, there is
no such thing as α-opic lx, melanopic lx or rhodopic lx (etc.).
4 Conclusions
The success of international standardized metrologies depends on them being both
clear and accurate enough that they can be adopted in as wide a variety of applica-
tions as possible. This depends on achieving sufficient consistency, between systems
and over time, whilst reflecting the underlying phenomenon faithfully. Photometric
and colorimetric standards are an average between individuals, which means
that interindividual variation is particularly important amongst the factors which
limit the need to produce an endless stream of decimal places. Another factor is
measurement uncertainty—photometry depends on the uncertainty of the radiome-

try, combined with the uncertainty in the psychophysical or neurophysiological
response. This is often an important limitation in work related to IIL responses.
Cone fundamentals and IIL responses are active research areas that will surely be
explored further, and that may lead to improved descriptions and new applications.
There are known shortcomings with established metrics, but continuously adopting
research findings could be significantly disruptive to industrial applications. On the
other hand, without updates at the level of international standards, innovation in light-
ing design, public health messages and trade in new technologies could be stifled.
Lastly, it would be remiss not to acknowledge that there exist assorted alternative
national and independent recommendations on IIL responses, particularly for build-
ing and lighting design. The distinction between scientific theory and industry
norms is sometimes unclear. This situation brings the risk of leaving people bewil-
dered and unsure who or what to believe. Where possible, subject experts should be
encouraged to participate in evidence-based consensus approaches, and researchers
should be encouraged to utilize the resulting standards and recommendations
whenever possible in testing models and theories.
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CIE, 2018c. CIE Alpha-Opic Toolbox. CIE, Vienna.
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CIE, 2019b. ISO/CIE 11664-1:2019 Colorimetry—Part 1: CIE Standard Colorimetric
Observers and Mesopic. CIE, Vienna.
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Figueiro, M.G., Gamlin, P.D., Lockley, S.W., O’Hagan, J.B., Price, L.L.A.,
Provencio, I., Skene, D.J., Brainard, G.C., 2014. Measuring and using light in the mela-
nopsin age. Trends Neurosci. 37 (1), 1–9.
Provencio, I., Jiang, G., Willem, J., Hayes, W.P., Rollag, M.D., 1998. Melanopsin: an opsin in
melanophores, brain, and eye. Proc. Natl. Acad. Sci. 95, 340–345.
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wavelength cones. Vision Res. 39, 2901–2927.
CHAPTER
Optical stimulation
systems for studying
human vision
Pablo A. Barrionuevoa,*, Oscar U. Preciadoa,b,
2
Marı́a L. Sandoval Salinasa,c, and Luis A. Issolioa,b
a
Instituto de Investigación en Luz, Ambiente y Visión, Consejo Nacional de Investigaciones
Cientı´ficas y T
ecnicas—Universidad Nacional de Tucumán, Tucumán, Argentina
b
Departamento de Luminotecnia, Luz y Visión, Facultad de Ciencias Exactas y Tecnologı´a,
Universidad Nacional de Tucumán, Tucumán, Argentina
c
Instituto de Investigaciones en Biodiversidad Argentina, Facultad de Ciencias Naturales e
Instituto Miguel Lillo, Universidad Nacional de Tucumán, Tucumán, Argentina
*Corresponding author: e-mail address: pbarrionuevo@herrera.unt.edu.ar
Abstract
This chapter describes the most common setups that scientists use for generating light
stimulation, from lab-made approaches to commercially available technologies. The studied
optical stimulation systems are divided into nonimage-forming and image-forming arrange-
ments. Two classical systems widely used are among the first: the Maxwellian view system
and the Ganzfeld stimulator. Between the image-forming arrangements, the focus is on
approaches that consider off-the-shelf devices and the recent appearance of multi-primary
displays, which allow the inclusion of more primaries and the generation of stimulation for
independent and combined photoreceptor and postreceptoral excitations. Some of the several
limitations that can have important implications in research practice are also examined, such as
those related to color gamut, sampling frequency, light range, and spatial resolution.
Since experimentation on how optical radiation is processed by the human neural system
requires the reliability of the parameters and variables under study to be assured, the charac-
terization and consequent calibration of experimental devices are essential. Therefore the chap-
ter discusses a set of characterization and calibration principles that researchers should consider
when carrying out experiments with the described optical stimulators. Outstanding characteris-
tics are stimulator response curve, primaries’ spectral power distribution, additivity, modulation
transfer function, and temporal stability. Finally, some possible sources of artifacts that re-
searchers should consider when these stimulators are used are presented. Throughout this last
section, data based on different optical stimulator measurements is provided.
Keywords
Calibration principles, Display limitations, Ganzfeld, Maxwellian view, Monitors,
Multi-primary systems, Optical arrangements, Physical characterization, Projectors, Silent
substitution
Progress in Brain Research, Volume 273, ISSN 0079-6123, https://doi.org/10.1016/bs.pbr.2022.04.003
13
14 CHAPTER 2 Optical stimulation systems for studying human vision
Abbreviations
CRT Cathode ray tube
DLP Digital light processing
DMD Digital micromirror device
ERG Electroretinography
Fs Sampling frequency
ipRGC intrinsically photosensitive retinal ganglion cell
LC Liquid crystal
LCD Liquid crystal display
LCoS Liquid crystal on silicon
LED Light emitting diode
MTF Modulation transfer function
OLED Organic light emitting diode
PSF Point spread function
SPD Spectral power distribution
1 Introduction
Experimentation in humans to study how optical radiation is processed by the
neural system requires physical characterization and careful control of the radiant
stimulation. Researchers have employed different optical configurations over the
years, depending on the attribute they intended to study and the level of detail they
needed to achieve, but also depending on the technology available. The use of per-
sonal computers together with different display technologies and lab-made optical
arrangements has facilitated the development of this field, although it requires char-
acterization and calibration processes to guarantee the reliability of the parameters
and variables under study.
In this chapter, we describe the most common technologies that scientists cur-
rently use for generating light stimulation, from lab-made approaches to commer-
cially available technologies. We also introduce characterization and calibration
principles that researchers should consider when carrying out experiments with
the described optical stimulators. These arrangements for optical stimulation are
common nowadays in vision research laboratories. Since complex light stimulation
can be useful for studying the circadian system (Spitschan, 2021), chronobiologists
or sleep researchers could also be interested in the arrangements discussed.
To start, we briefly explain silent substitution, which is a method widely used
in the arrangements described in the following sections.
1.1 The silent substitution method
The electrical response of a photoreceptor results from the integration of the total
light quanta absorbed in its spectral sensitivity wavelength range. Therefore this
response is not dependent on the light wavelength. This is called the principle of
univariance (Mitchell and Rushton, 1971). This photoreceptor property, together
with the overlapping nature of photoreceptor spectral sensitivities, has led to the
2 Optical stimulation systems 15
development of the silent substitution technique (Estevez and Spekreijse, 1982).

With this technique it is possible to effectively obtain independent photoreceptor-
driven responses while maintaining the same adaptation for all photoreceptors.
Evidence of silent substitution usage comes from the beginning of the 20th century
(Ishihara, 1906). In this technique, the radiances of n lights are modulated to silence
up to n 1 photoreceptor types, while the nonsilenced photoreceptor type received
controlled contrast stimulation (Pokorny and Cao, 2010; Spitschan and Woelders,
2018). To apply the silent substitution method, the stimulus needs to be composed
of independent spectrally different light sources (usually light-emitting diodes, or
LEDs). For example, in the mesopic region where rods and S-, M-, and L-cones
are active (Zele and Cao, 2015), the system should have at least four types of LEDs.
2 Optical stimulation systems

2.1 Nonimage-forming arrangements
Classical optical arrangements used incandescent and arc lamps for retinal stimula-
tion; light intensity was controlled using filters, beam splitters, shutters, rotating
discs, etc. Therefore, for setting up vision experiments, the researcher needed to have
good training in optics (Pokorny and Smith, 2020). With the incorporation of LEDs,
their associated electronics, RGB displays, and computation tools into research
laboratories, it was easier to develop complex stimulation to investigate vision at
different domains with relatively basic knowledge of optics. Here we describe
two classical nonimage-forming arrangements that are widely used even now by
researchers: the Maxwellian view system and the Ganzfeld stimulator.
2.1.1 Maxwellian view systems

From an optical point of view, regular displays—such as monitors and projectors—
are characterized to allow free or “Newtonian” vision, where the image is formed on
the retinal plane by the optics of the eye (Fig. 1A). However, with these devices, it
is not possible to be certain about the light reaching the retina or the illuminated
retinal area (Westheimer, 1966). An alternative optical approach was introduced
by Maxwell (1860). In his arrangement, the image is placed at the pupil plane
(Fig. 1B). The observer sees the lens uniformly filled with light and the stimulation
area is determined by the lens size (Wyszecki and Stiles, 2000). The advantages of
this arrangement are: (1) it can produce stimulus where the focus, shape, and size are
independent of the retinal illuminance; (2) the light levels reaching the retina can be
higher than those produced by free-viewing systems (Burns and Webb, 2010); and
(3) the retinal image is free of intraocular scattering generated by the iris and the
sclera (van den Berg, 1995). One disadvantage of this arrangement is that it is
not adequate for very small light sources (Wyszecki and Stiles, 2000). Also, the
observer’s eye has to be located at the correct position during the entire experimental
session to avoid any artifacts caused by unwanted retinal illuminance changes. To
satisfy this condition, many experimenters used a bite bar with the observer’s dental
impressions together with an XYZ positioning system (Matesanz et al., 2011).
FIG. 1
Optical arrangements. (A) In a Newtonian or free-viewing arrangement, images are formed on the retinal plane by the optics of the eye. (B) In a
Maxwellian view arrangement, images are placed at the pupil plane, and on the retinal plane images correspond to that of the lens
uniformly filled with light. (C) Schematic representation of a Ganzfeld stimulator, which uniformly stimulates vast portions of the retina.
A minimal optical arrangement to obtain Maxwellian vision is shown in

Fig. 1B. A small light source, such as an LED, is imaged on the plane of the subject’s
pupil by a biconvex lens. This lens should be achromatic to reduce optical aberrations
and avoid breakup of primaries mixing when several monochromatic lights are used.
The observer’s eye focuses on the lens or an object next to it. The lens is seen
uniformly illuminated. Usually, an artificial pupil smaller than the minimal natural
pupil is located next to the eye to control retinal illuminance. If the image of the light
source is smaller than the minimal pupil aperture, the artificial pupil or the topical
mydriatics for pupil dilation are not necessary.
A potential problem with this arrangement is that the eye has to accommodate to
focus on the lens. A small change in the position of the object will produce a defocus
on the image. To overcome this issue, more complex arrangements were proposed
(Burns and Webb, 2010); however, for many nonimage-forming experiments, the
setup of Fig. 1B is adequate.
Nowadays, these systems are still used, for example, to study retinal light adap-
tation (Barrionuevo et al., 2018a; Gloriani et al., 2016). Maxwellian view arrange-
ments provide a way to avoid unwanted effects of pupil-generated intraocular light
modulation and, therefore, illuminance reaching the retina can be easily controlled.
An experimental setup using the Maxwellian view consists of the multi-primary
photostimulators, which are addressed in the next section.
2.1.1.1 Multi-primary photostimulators

First introduced by Pokorny and Smith’s laboratory (Pokorny et al., 2004; Sun et al.,
2001a), a four-primary photostimulator allowed independent control of the stimulation
of rods and L-, M-, and S-cones using the silent substitution method (described in
Section 1). This methodology had definite advantages: to assess photoreceptor func-
tioning at the same adaptation level, retinal locus, and chromaticity; and to control
photoreceptor intrusion to postreceptoral pathways (Pokorny and Cao, 2010). The
four-primary photostimulator contained two optical branches, one used to produce a
central field and the second branch to illuminate a surrounding field. It has been used
in psychophysical and electrophysiological settings to study brightness induction (Sun
et al., 2001b), color perception (Cao et al., 2005), chromatic discrimination (for exam-
ple, Cao et al., 2008), flicker perception (for example, Cao et al., 2006), photoreceptor
temporal contrast sensitivity (Huchzermeyer and Kremers, 2017; Sun et al., 2001a),
temporal processing (for example, Cao et al., 2007), rod-cone interaction (for a review
see Buck, 2014), and the early stage of visual processing (for example, Cao et al.,
2010), among other subjects. A five-primary photostimulator producing high photopic
light levels (>2000Td) has been developed for the study of the melanopsin activation
in intrinsically photosensitive retinal ganglion cells or ipRGCs (Cao et al., 2015).
With this device, the contribution of melanopsin to pupil responses (Barrionuevo
and Cao, 2016) and visual processing (Zele et al., 2018, 2019) was studied. The high
light level produced and the temporal and spectral control makes the five-primary
photostimulator suitable for circadian research.
For a deeper explanation of Maxwellian view arrangements and optical compu-
tation, we encourage the reader to consult the works of Burns and Webb (2010),
Westheimer (1966), and Wyszecki and Stiles (2000).
2.1.2 Ganzfeld stimulators

Ganzfeld (German for “whole field”) or full-field stimulators are used to uniformly
stimulate a vast portion of the retina. This stimulator is inspired by the integrating
sphere used in colorimetric practice (Wyszecki and Stiles, 2000). A full-field system
consists of a semispherical dome illuminated in such a way that the observer only
receives reflected but not direct light from the light source (Fig. 1C). The interior
of these domes ideally consists of a perfectly diffusive (Lambertian) spectrally
nonselective white surface. On a Lambertian surface, the luminous intensity in
any direction varies with the angle cosine between that direction and the normal
to the surface (Wyszecki and Stiles, 2000).
The color of the stimulation is given by the light source. In old commercial
devices, the illuminant consists of incandescent lamps for uniform illumination
and a xenon gas discharge lamp for flashes. Modern devices contain LED illuminants
of different colors, which is convenient for chromatic control. The observer sees a
uniform and unstructured illuminated field (Hogg, 2006), and the system usually
includes one or several fixation spots. Also, these systems can include a camera
for pupillary recordings (Barrionuevo et al., 2014; Tripolone et al., 2019).
The following section briefly explains some of the main uses of this type
of device.
2.1.2.1 Electrophysiology
These systems are used in electroretinography (ERG), where a full-field stimulation
is needed to produce a significant electrical response of the retina. In electrophysi-
ology, full-field stimulation provides information of the peripheral retina and enables
the differentiation between outer and inner retina function following different testing
protocols according to the stimulus and state of adaptation (McCulloch et al., 2015;
Robson et al., 2018). New protocols including chromatic stimulation have been
proposed to discriminate between photoreceptors: for example, S-cone responses
(Perlman et al., 2020), melanopsin responses (Rabin et al., 2020), and mesopic
rod-cone responses (Barrionuevo et al., 2018b).
2.1.2.2 Pupillometry
Ganzfelds are being widely employed in chromatic pupillometry. This technique
consists of the recording and parameterization of the pupillary light reflex under
chromatic stimulation, generally with red and blue stimuli. Full-field stimulation
is preferred since the pupil seems to integrate responses over larger areas than
image-forming vision (Lei et al., 2014; Park and McAnany, 2015), and therefore
the pupillary response is much higher with a more extensive stimulation area. It is
possible to generate chromatic stimuli that enhance pupillary responses driven by
activation of cones, rods, and ipRGCs, and therefore to detect anomalies in retinal
function. The technique has reached such maturity that protocols for clinical studies
were proposed (Kelbsch et al., 2019; Rukmini et al., 2019). Full-field stimulators
were used in chromatic pupillometry with promising results to detect pathologies
such as retinitis pigmentosa (Kardon et al., 2011), glaucoma (Duque-Chica et al.,
2018), diabetes (Feigl et al., 2012), and Leber congenital amaurosis (Kawasaki
et al., 2012).
2.1.2.3 Silent substitution in Ganzfelds

A more sophisticated technique with full-field stimulation to study the pupil light
reflex is the silent substitution method (see Section 1). Applying this technique in
full-field commercial stimulation devices, it was possible to study, for example,
rod-cone interaction (Barrionuevo et al., 2014), the M-cone contribution (Murray
et al., 2018), and the nonlinearities in the pupillary neural circuit (Barrionuevo
et al., 2018b). Also, lab-made full-field arrangements were used in conjunction with
silent substitution to study the contribution of ipRGCs in steady-state pupil response
(Tsujimura et al., 2010; Vienot et al., 2010).
2.2 Image-forming arrangements

2.2.1 Monitors
For a long time, to carefully present a visual stimulus, the most prominent technology
used by researchers was the cathode ray tube (CRT) monitor. Most off-the-shelf CRT
monitors provided several advantages for a vision laboratory: great color gamut,
fast response, good spatial resolution, self-illumination, easy control of the RGB
primaries, and long lifetime with tolerable depreciation. However, the CRTs were
discontinued by the intrusion of new technologies that offered flat displays, a major
advantage compared to the bulkiness of CRTs, which were rapidly preferred by the
general public. Therefore researchers started assessing the suitability of new displays
for research purposes. Primarily, many works focused on comparing the performance
of CRTs with liquid crystal display (LCD) monitors, plasma devices, and organic
light-emitting diode (OLED) displays. Nowadays, CRT technology is obsolete and
monitors of this kind are a rare find for consumer purposes. Although the CRT
monitors are still being used in vision laboratories, we briefly describe here LCD
and OLED monitors available on the market, and some configurations have
been shown mature enough to be suitable for research purposes (Ito et al., 2013;
Zhang et al., 2018). We encourage the readers with interest in CRTs to look into these
reviews: Wheeler and Clark (1992), Brainard et al. (2002), and Bauer (2015).
2.2.1.1 Liquid crystal display

A transmissive LCD generally consists of a backlight and a diffuser, followed by
the liquid crystal (LC) with an array of transistors sandwiched by two polarizers
(Lee et al., 2020). Light coming from the backlight—which is always turned
on—is polarized when it enters the LC and each transistor of the array can vary
the direction of polarization controlling the applied voltage to the LC. Then the po-
larized light passes through a second polarizer, which is 90 degrees axially rotated
with respect to the first polarizer. If there is no variation in the LC of the polarization
direction, the light is not passing and the observer sees a dark pixel. On the contrary,
if the direction of polarization is varied 90 degrees by the LC, light is passing through
and the observer sees a bright pixel. If the LC varies the direction angle between
0 and 90 degrees, gray light levels are achieved.
2.2.1.2 Organic light-emitting diode

OLED displays are structurally formed by several organic material layers between
two conductive arrays called the anode and cathode. In the organic material, the
recombination of an electron with a hole (lack of an electron in an atom), liberates
a photon when a current source is connected to the anode and cathode terminals;
this phenomenon is called electroluminescence and constitutes the functional basis
of OLED displays (Tsujimura, 2017). Cathode and anode terminals are perpen-
dicularly arranged to form a matrix; each intersection in this matrix is therefore
an independent source of light. By modulating the electrical current, different
luminosities can be achieved.
2.2.2 Projectors
In the last decades, projectors have gained portability, efficiency, and image quality,
being a good option to generate visual stimulation in a laboratory setting. Although
there are research-dedicated devices, for many experimental needs an off-the-shelf
projector is enough. A projector is usually composed of a high-intensity light source,
the image generator or imager, and optics to project the image onto the screen.
Here we describe these components, in order to understand the functioning of this
technology.
2.2.2.1 Light source

Typically, the light source used in projectors is a gas discharge lamp, for example, a
xenon lamp or a high-pressure lamp. Discharge lamps have a broad spectrum and,
through different filters or dichroic mirrors, it can be split into the RGB primaries.
In recent years, lasers are increasingly being used in projectors as light sources. The
laser usually emits in the short-wavelength range (B primary) and through a
phosphor wheel or LEDs, the R and G primaries are obtained. Both light sources
accomplish two important criteria: high brightness and high efficiency (Lee et al.,
2020). Laser-based projectors are more efficient than discharge lamp-based projec-
tors, produce greater color gamut and brightness, have a longer lifetime, and are
mercury-free; however, they are more expensive. Furthermore, the image can be
affected by the speckle noise due to the coherent nature of the lasers (Akram and
Chen, 2016), which is observed as a grainy pattern superposed on the projection
(Pauwels and Verschaffelt, 2017).
2.2.2.2 Imager
Projectors using LCD technology generally include three transmissive LCD panels.
Light from the source is divided by dichroic mirrors into three components corre-
sponding to the RGB primaries, each primary light is processed accordingly by
one LCD panel, and the images generated by the three panels are combined in a
full-color image using a prism (Fig. 2A).
The second kind of LCD-based projector is called a liquid crystal on silicon
(LCoS) system. In LCoS projectors, light is reflected instead of being transmitted.
Light enters the LC layer, which is behind a polarizer and a color filter, and then
the light is reflected on a mirror layer. The LC polarization is commanded by a
CMOS layer behind the mirror (Yang and Wu, 2014).
On the other hand, in digital light processing (DLP) projectors (Fig. 2B), the
image generator is a digital micromirror device (DMD). The DMD is an arrangement
of microscopic mirrors. Each mirror can be tilted between two orientations for
reflecting light to the projecting optics. The different gray levels are achieved when
the mirror is moved at high velocity and the relative time in each position is mod-
ulated; this is called pulse width modulation (Florence and Yoder, 1996). To produce
a colorful image, these projectors contain a spinning color-filter wheel sectioned
with at least three primaries. Light passes through a portion of the wheel and digital
information for each primary is synchronized with the angular position of the wheel.
Since the wheel is rotating at a velocity higher than the temporal fusion frequency of
the eye, color additivity is produced by temporal fusion.
2.2.2.3 Projection optics

Light from the small pixels of the imager needs to be projected on a distal screen. For
this purpose, an arrangement of lenses is used. Common problems associated with
the optics used in projectors are low efficiency and poor uniformity. Furthermore,
this arrangement needs to be relatively small to coexist with other optics from
the light path arrangement, but if the imager size increases, the optics increase in
proportion (Chang and Shieh, 2000).
2.2.2.4 Screens
Projection screens are divided into two broad groups, front and rear projecting
screens. Rear projecting screens are transmissive, so part of the light is absorbed
or reflected, which, in general, produces an important reduction of brightness com-
pared to front projecting screens, where the most part of the light is reflected with
small loss by reflection. Also, a rear projecting system needs a room with larger
dimensions. One desirable feature for photometric and radiometric purposes is that
the screen surface has Lambertian diffusion.
FIG. 2
Schematics of optical treatment and components in a top view of (A) LCD and (B) DLP
projectors. (C) Optical arrangement of a multi-primary display. The images of two monitors are
overlapped and using spectral filters the spectral composition of the green primaries is
modified in order to have four primaries at each pixel (the representation is based on the
proposal of Cao’s laboratory).
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the type used with post cards is employed.—Thomas R. Milligan,
Fort Worth, Texas.
Three-Caster Truck for Moving Crates and
Furniture
A convenient truck for handling heavy objects, especially in the
home where commercial devices for this purpose are not available,
is shown in the illustration. It consists of a frame built up of three 1¹⁄₄
by 2 by 14-in. strips, fixed to a disk, ⁷⁄₈ by 12 in. in size. Revolving
casters are mounted under the ends of the arms, giving great
freedom of movement in transporting loads. The three-caster
arrangement is better than the use of four casters, because it
accommodates itself to irregularities in the floor.—Armydas E.
Sturdivant, Muncie, Ind.
¶The burner of an acetylene bicycle lamp, fitted to an ordinary gas

jet, will produce a very hot flame.
Inserting or Correcting on Typewritten Bound
Sheets
It is frequently necessary that typewritten insertions or corrections
be made in papers which have been typed and bound, usually by
rivets along the top edge. It is difficult to remove the rivets and
replace them satisfactorily. To make such insertions, feed a blank
sheet of paper into the typewriter until its feeding edge is even with
the upper edge of the guide fingers. Then insert the bottom edge of
the sheet on which the correction is to be made, underneath the
sheet already in the machine; reverse-feed the sheet to be corrected
into the typewriter. Corrections may then be made in the usual
manner.—A. J. Cook, Pittsburgh, Pa.
Tire Pump Made of Gas Piping
A tire pump actuated by the explosions in the cylinder of an
automobile engine, and made of materials easily obtainable, is
shown in detail in the illustration. A section of iron pipe of a diameter
to fit one of the ports, and about 4 in. long, as shown in Fig. 2, is
provided with a reducer, a tee, and two caps. The inside of the pipe
is turned up true in a lathe and fitted with a piston, the head for which
is shown in Fig. 1. The head is built up of a wooden drum, washers,
and leather packing disks. The washers should be slightly smaller in
diameter than the chamber, and the leather disks make it air-tight.
Fig. 1
Fig. 2
Fig. 3
Automobile Tires may be Inflated Quickly by the Use of This Homemade

Pump
The tee is provided with two valves, as shown in the detail
sectional view, Fig. 3, the upper ball valve retaining the pressure in
the pipe line, and the other valve admitting air above the piston. The
ball valve has a seat of lead, and a ribbed fitting is fixed to the pipe
cap above it, to provide a fastening for the hose. The cap of the
other valve is fitted with a spring, adjustable on a threaded pin.
A light spring fixed to the upper side of the piston, as shown in Fig.
2, forces the piston back to its lower position after the force of the
explosion in the cylinder has acted upon it. The cylinder of the pump
should be oiled, and a reinforced rubber hose should be used to
conduct the air under pressure to the tire.
The operation is simple: The initial stroke of the engine cylinder
draws the piston down, bringing in a charge of air from the valve
ports at the side of the tee; the following compression and explosion
strokes drive it back, forcing the air out of the ball valve, as the other
valve closes on the upstroke of the piston. A large tire may be
pumped up with this device in from five to ten minutes.
Aid in Ruling Uniform Cards or Sheets
When a number of cards or sheets are to be ruled with either
horizontal or vertical divisions, the following method will be found a
timesaver: Rule one of the cards as a sample. Place it on the
drawing board with its lower edge set against two thumb tacks driven
part way in. Rule extensions of the lines on the card to a sheet of
paper fastened to the board under the card. Set a third thumb tack at
the right edge of the card as a guide. By placing the cards to be
ruled against the three thumb tacks in the position of the original
card, the rulings may be made quickly with the marks on the paper
backing as guides.—W. P. Shaw, Stratford, Can.
Tinned Staples for Bell-Circuit Wiring
The Strip is Cut into Squares Which are Folded and Driven into the Support,
Binding the Wire
Inexpensive and practical staples for binding wires used in bell

circuits, or for similar purposes, may be made from a strip of tinned
sheet metal, about ¹⁄₂ in. wide. The strip is cut on the sides, as
indicated in the sketch, and the squares are broken off as needed.
They may be bent quickly to the shape shown, and are used by
placing them over the wire and driving the points into the wall or
other support. If properly made, they will not injure the covering of
the wire and are more satisfactory than wire staples.

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Circadian and Visual Neuroscience -

Circadian and Visual

Tanya Calvey, Johannesburg, South Africa

Circadian and Visual

First edition 2022

Copyright © 2022 Elsevier B.V. All rights reserved.

For information on all Elsevier publications

Publisher: Zoe Kruze

CHAPTER 1 Circadian and visual photometry.................................1

CHAPTER 2 Optical stimulation systems for studying human

CHAPTER 3 Hyperspectral characterization of natural

2.1 Capturing images of a mirror sphere .................................... 40

CHAPTER 4 Endogenous functioning and light response of the

CHAPTER 5 Light-dependent effects on mood: Mechanistic

CHAPTER 6 Rodent models in translational circadian

CHAPTER 7 Slow vision: Measuring melanopsin-mediated

CHAPTER 8 Beyond irradiance: Visual signals influencing

3. Evidence for inner and outer retinal sources of circadian

CHAPTER 9 Circadian photoreception: The impact of light

CHAPTER 10 Modeling (circadian)................................................181

CHAPTER 11 Visual encoding: Principles and software..............199

CHAPTER 12 Visual psychophysics: Luminance and color..........231

CHAPTER 13 Aging of visual mechanisms....................................257

3. Senescent changes in sensitivity of rod and cone pathways ..... 262

CHAPTER 14 Seeing and sensing temporal variations in

CHAPTER 15 Light in ecological settings: Entrainment,

3.2 Position within a time zone ................................................ 313

Circadian and visual

Progress in Brain Research, Volume 273, ISSN 0079-6123, https://doi.org/10.1016/bs.pbr.2022.02.014

1 The basis of physical photometry

K m ¼ K cd =V ðλcd Þ ¼ 683:002 lm=W (2)

K 0 m ¼ Kcd =V 0 ðλcd Þ ¼ 1,700.05 lm=W (3)

1750 1700.048 lm/W Photopic

750 683.002 lm/W

1=γ x ¼ max ½ sx ðλÞ h f (7)

1.1 Additional notes to Eqs. (1) to (7)

Having specified (x, y), z becomes redundant:

10° cone fundamentals

3.1 Additional notes to Eqs. (14) and (15)

measurement uncertainty—photometry depends on the uncertainty of the radiome-

CIE, 2018b. CIE 015:2018 Colorimetry, fourth ed. CIE, Vienna.

development of the silent substitution technique (Estevez and Spekreijse, 1982).

2 Optical stimulation systems

2.1.1 Maxwellian view systems

A minimal optical arrangement to obtain Maxwellian vision is shown in

2.1.1.1 Multi-primary photostimulators

2.1.2 Ganzfeld stimulators

2.1.2.3 Silent substitution in Ganzfelds

2.2 Image-forming arrangements

2.2.1.1 Liquid crystal display

2.2.1.2 Organic light-emitting diode

2.2.2.1 Light source

2.2.2.3 Projection optics

Folders Like That in Figure 1 may be Glued to Form Envelopes, Figure 2, or

I have derived considerable satisfaction from the use of a yardstick

The uppers from a pair of worn-out shoes may be made into a

Scenery for an Amateur Theatrical Performance was Made by Painting

Scenery for a home-talent theatrical performance was required,

¶The burner of an acetylene bicycle lamp, fitted to an ordinary gas

Automobile Tires may be Inflated Quickly by the Use of This Homemade

Inexpensive and practical staples for binding wires used in bell

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