- •An Organ of Exquisite Perfection
- •Optical Path
- •Retinal Photoreception
- •Photoreception Optics
- •Photoreception Biochemistry
- •Membrane Voltages
- •Blind Spot
- •Retinal Pathways
- •Through Pathway
- •Receptive Fields
- •Lateral Pathway
- •Retinal Ganglion Cells
- •Retinal Glia
- •References
- •Development of the Foveal Specialization
- •Introduction
- •Foveal Development
- •Specification of Foveal Location
- •Formation of a Rod-Free Zone
- •Cones, Ganglion Cells, and Initial Pit Formation
- •Deep Foveal Pit Formation
- •Foveal Hypoplasia
- •Conclusions and Perspectives
- •Acknowledgments
- •References
- •An Update on the Regulation of Rod Photoreceptor Development
- •Introduction
- •Brief Overview of Retinal Development and Early Stages of Rod Photoreceptor Differentiation
- •Transcription Factors
- •Basic Helix-Loop-Helix Genes
- •Nuclear Receptors
- •Retinoic Acid/Retinoic Acid Receptors
- •Wnt/Frizzled Pathway
- •Taurine
- •Ciliary Neurotrophic Factor/Leukemia Inhibitory Factor/Pleiotrophin/Signal Transducer and Activators of Transcription 3/SOCS
- •Conclusions and Future Prospects
- •References
- •Introduction
- •Retinal Adhesion
- •Physiology of Retinal Adhesion
- •Molecular Mechanisms of Retinal Adhesion
- •Significance of Retinal Adhesion for Retinal Function
- •Photoreceptor Outer Segment Renewal
- •Physiology of Outer Segment Disk Assembly and Disk Shedding
- •Physiology of RPE Engulfment of Shed Outer Segment Fragments
- •Molecular Mechanisms of Shedding and RPE Phagocytosis
- •Significance of Photoreceptor Outer Segment Renewal for Retinal Function
- •Perspective
- •Acknowledgments
- •References
- •Molecular Biology of IRBP and Its Role in the Visual Cycle
- •Introduction
- •IRBP Protein Studies
- •IRBP Null Mice
- •IRBP Induces Experimental Autoimmune Uveitis
- •IRBP Expression During Development
- •Variability in IRBP Expression
- •Molecular Biology of IRBP
- •IRBP Genomic Cloning
- •Evolution of IRBP
- •Identification of DNA cis-Acting Controlling Elements: In Vitro and In Vivo Experiments
- •Transcription Factors and their Role in the Control of IRBP Expression
- •Rx/rax Transcription Factor
- •NrL Transcription Factor
- •Crx Transcription Factor
- •OTX2 Transcription Factor
- •Transgenic Mice
- •Repressors of IRBP Gene Expression
- •Summary and Conjecture
- •Acknowledgments
- •References
- •Regulation of Photoresponses by Phosphorylation
- •Introduction
- •Cone-Specific Kinase, GRK7
- •Protein Kinase C
- •Cyclin-Dependent Kinase
- •Tyrosine Kinases
- •Protein Phosphatases
- •Conclusion
- •References
- •The cGMP Signaling Pathway in Retinal Photoreceptors and the Central Role of Photoreceptor Phosphodiesterase (PDE6)
- •Regulation of Intracellular cGMP Levels in Photoreceptor Cells
- •Downstream Targets of cGMP Action in Photoreceptor Cells
- •cGMP-Dependent Protein Kinase
- •Cyclic Nucleotide-Gated Ion Channels
- •PDE6 Is a High-Affinity cGMP-Binding Protein
- •Compartmentation of cGMP Signaling in Photoreceptor Outer Segments
- •Physiology of the Photoreceptor Response to Light
- •Biochemical Cascade of Visual Excitation
- •Central Components of the cGMP Signaling Pathway
- •Termination and Adaptation of the Light Response
- •Deactivation of Rhodopsin
- •Deactivation of Transducin
- •Deactivation of PDE6
- •Activation of GC
- •Regulation of the CNG Ion Channel
- •Photoreceptor PDE (PDE6) Structure and Function
- •The Cyclic Nucleotide Phosphodiesterase Superfamily
- •Subunit Composition of Rod and Cone PDE6 Holoenzyme
- •Catalytic Subunit
- •Regulatory GAF Domain
- •Catalytic Domain
- •C-Terminal Prenylation
- •PDE6 Has Evolved to Meet the Special Demands of the Central Effector of Visual Transduction
- •PDE6 Regulation
- •Transducin Activation of Rod PDE6 During Visual Excitation
- •Functions of the Regulatory cGMP-Binding GAF Domains of PDE6
- •Potential PDE6 Regulatory Binding Proteins
- •Glutamic Acid-Rich Protein 2
- •Conclusions
- •Acknowledgments
- •References
- •Rhodopsin Structure, Function, and Involvement in Retinitis Pigmentosa
- •Introduction
- •Historical Perspective
- •Rhodopsin, Localization, and Signaling
- •Dark State and Activation
- •Structural Analysis
- •Electron Cryomicroscopy and Crystal Structure
- •Nuclear Magnetic Resonance
- •Cysteine Mutagenesis and Electron Paramagnetic Resonance
- •Other Approaches
- •Retinitis Pigmentosa
- •Transmembrane RP Rhodopsin Mutants
- •Cytoplasmic RP Rhodopsin Mutants
- •Intradiskal RP Rhodopsin Mutants
- •Implications of Receptor Misfolding
- •Nongenetic Contributions to RP
- •Conclusion
- •References
- •Multiple Signaling Pathways Govern Calcium Homeostasis in Photoreceptor Inner Segments
- •Introduction
- •Overview of Ca2+ Regulation in the Inner Segment
- •Voltage-Operated Calcium Channels Play a Central Role in Inner Segment Calcium Regulation
- •Ca2+ Channels in Rods and Cones
- •Photoreceptor Malfunction and Degeneration
- •Therapeutic Strategies
- •Development
- •Acknowledgments
- •References
- •The Transduction Channels of Rod and Cone Photoreceptors
- •The Role of CNG Channels in Photoreceptor Physiology
- •The Activation Phase of the Light Response
- •Recovery After a Light Stimulus and Adaptation to Continuous Illumination
- •CNG Channels in the Synaptic Transmission of Cone Photoreceptors
- •The Molecular Composition of CNG Channels
- •The Basic Activation Properties of CNG Channels
- •Transmembrane Topology and Functional Domains
- •The Cyclic-Nucleotide-Binding Domain
- •The Amino Terminal Domain and Modulation by Calmodulin
- •The P Region
- •The GARP Domain of CNGB1
- •Modulation by Phosphorylation and All-trans Retinal
- •Synthesis, Maturation, and Targeting of CNG Channels
- •Visual Dysfunction Caused by Mutant CNG Channel Genes
- •References
- •Appendix
- •Visual Dysfunction Caused by Mutant CNG Channel Genes
- •Mutations in CNGA1 and CNGB1 Associated with Retinitis Pigmentosa
- •Mutations in CNGA3 and CNGB3 Associated with Cone Dysfunction
- •References
- •Rhodopsins in Drosophila Color Vision
- •Introduction
- •Anatomy and Molecular Aspects of Color-Sensitive Opsins in the Drosophila Eye
- •Structure of the Drosophila Eye: Ommatidia, Photoreceptors, and Rhodopsins
- •Molecular Genetics and Evolution of Rh5 and Rh6
- •Development and Patterning of Rhodopsins for Drosophila Color Vision
- •Mutually Exclusive Rhodopsin Expression
- •Transcription Factors Specify Outer from Inner Photoreceptors and Distinguish R7 from R8
- •A Stochastic Decision Induces Rhodopsins in R7 Photoreceptor
- •A Bistable Feedback Loop Specifies R8 Photoreceptor Subtype and Expression of Rh5 and Rh6
- •Comparison Between Mammalian and Drosophila Color Vision Rhodopsins
- •Human Color-Sensitive Opsins
- •Conclusion
- •References
- •INAD Signaling Complex of Drosophila Photoreceptors
- •Introduction
- •Identification of the INAD Signaling Complex
- •Function of the INAD Signaling Complex
- •Information Transfer From Rhodopsin to the Signaling Complex BY the Visual G Protein
- •Signaling Complexes in Vertebrate Photoreceptor Cells
- •Acknowledgments
- •References
- •Visual Signal Processing in the Inner Retina
- •Introduction
- •Visual Information is First Processed in the OPL
- •Bipolar Cells form Parallel Pathways and Provide Excitatory Input to the IPL
- •Functional Stratification of the IPL
- •ON and OFF Response Stratification
- •Sustained and Transient Response Stratification
- •Synaptic Mechanisms Shape Excitatory Signals in the IPL
- •Glutamate Release Is Tonic and Graded
- •Transporters Terminate Excitatory Signaling to Ganglion Cells
- •Postsynaptic Glutamate Receptor Properties Shape Ganglion Cell Excitation
- •Modulating Glutamate Release Shapes Excitatory Responses
- •Amacrine Cells Mediate Inhibition in the IPL
- •Presynaptic Inhibition
- •Asymmetric Presynaptic Inhibition
- •Presynaptic Inhibition Is Filtered by GABA Receptor Properties
- •Presynaptic Inhibition May Be Shaped by Transmitter Release Differences
- •Glycine, the Other Inhibitory Transmitter
- •Parallel Ganglion Cell Output Pathways
- •Ganglion Cells Encode Color Information
- •Directional-Selective Ganglion Cells
- •Intrinsically Photosensitive Ganglion Cells
- •Conclusions
- •References
- •Human Cone Spectral Sensitivities and Color Vision Deficiencies
- •Introduction
- •Overview
- •Transduction
- •Univariance, Monochromacy, Dichromacy, and Trichromacy
- •Trichromacy and Color-Matching Functions
- •Cone Spectral Sensitivities
- •Introduction
- •Cone Spectral Sensitivity Measurements
- •From Cone Spectral Sensitivities to Color-Matching Functions
- •Other Factors That Influence Spectral Sensitivity
- •Lens Pigment
- •Macular Pigment
- •Photopigment Optical Density
- •Changes with Eccentricity
- •Congenital Color Vision Deficiencies
- •Protan and Deutan Defects
- •Protanopia and Deuteranopia
- •Photopigment Variability and Protanomaly and Deuteranomaly
- •Tritanopia
- •Monochromacies
- •Cone Monochromacies
- •Rod Monochromacy
- •Conclusions
- •Acknowledgment
- •References
- •Luminous Efficiency Functions
- •Introduction
- •The Need for Luminous Efficiency
- •Psychophysical Measures of Luminous Efficiency
- •Factors that Influence Luminous Efficiency
- •Scotopic (Rod) Luminous Efficiency Function
- •Introduction
- •Univariance
- •International Standard
- •Photopic (Cone) Luminous Efficiency Function
- •Introduction
- •International Standards
- •Other Photopic (Nonadditive) Luminous Efficiency Functions
- •Mesopic (Rod-Cone) Luminous Efficiency Functions
- •Introduction
- •Models of Mesopic Luminous Efficiency
- •International Standard
- •Individual Differences Influencing Luminous Efficiency
- •Attenuation of Spectral Light by the Lens and Other Ocular Media
- •Attenuation of Spectral Light by the Macular Pigment
- •Optical Densities of the Photopigments
- •Relative Numbers of L and M Cones
- •Cone Pigment Polymorphisms
- •Directional Sensitivity
- •Variations in the Contribution of Chromatic Channels
- •Conclusions
- •References
- •Cone Pigments and Vision in the Mouse
- •Introduction
- •Prevalence and Spatial Distribution of Mouse Cones
- •Mouse Strain Variations
- •Mouse Cone Pigments
- •Cone Pigment Spectra
- •Evolution and Spectral Tuning of Mouse Cone Pigments
- •Regional Distribution of Mouse Cone Pigments
- •Expression of Mouse Cone Pigments
- •Cone Signal Pathways in the Mouse Retina
- •Cone-Based Vision in Mice
- •Assessment Techniques
- •Spectral Sensitivity
- •Spatial and Temporal Sensitivity
- •Color Vision
- •Targeted Deletions of Rods or Cones
- •Addition of New Cone Pigments
- •Mouse and Human Cone Vision
- •Acknowledgment
- •References
- •Multifocal Oscillatory Potentials of the Human Retina
- •Introduction
- •Recording Techniques
- •Underlying Mechanisms
- •The Influence of age and Gender
- •Disease-Related Changes
- •Origins of Single Potentials
- •Dichromats
- •Congenital Stationary Night Blindness
- •Topographical Alterations
- •Diabetes
- •Retinal Vessel Occlusion
- •Glaucoma
- •General Alterations
- •Vigabatrin Treatment
- •Conclusion
- •References
- •The Aging of the Retina
- •Introduction
- •Morphological Alterations
- •Neural Changes
- •Retinal Pigment Epithelium and Lipofuscin Formation
- •Bruch’s Membrane and Choroid
- •Retinal Function Changes
- •Age-Related Macular Disease
- •Conclusions
- •References
- •Aging of the Retinal Pigment Epithelium
- •Introduction
- •Aging Changes In the Fundus
- •Age-Related Changes In RPE Morphology
- •Melanosomes
- •Lipofuscin
- •Pigment Complexes
- •Mitochondria
- •Bruch’s Membrane
- •Functional Consequences of RPE Cell Aging
- •Phagocytic Load
- •The Effect of Lipofuscin on the RPE
- •Melanosomes
- •Antioxidant Capacity of the RPE
- •Lysosomal Enzyme Activity
- •Mitochondrial Damage in the RPE
- •Bruch’s Membrane Aging
- •Oxidative Stress and RPE Aging
- •The Relationship Between Aging and Retinal Pathologies
- •Summary and Conclusions
- •References
- •Visual Transduction and Age-Related Changes in Lipofuscin
- •Introduction: What is Lipofuscin?
- •Lipofuscin of the Retinal Pigment Epithelium
- •Composition of RPE Lipofuscin
- •Fluorescence Properties of RPE Lipofuscin
- •A2E as a Marker of Lipofuscin Accumulation
- •Factors Affecting Accumulation of RPE Lipofuscin
- •Phagocytosis and Autophagy
- •Role of Lysosomal Degradation
- •Role of Oxidative Stress
- •Role of Phototransduction in Accumulation of RPE Lipofuscin
- •Transient Buildup of All-trans Retinal in Photoreceptor Outer Segments as a Critical Factor for Lipofuscin Formation
- •Inhibition of the Retinoid Cycle Inhibits Lipofuscin Accumulation
- •Role of Exposure of the Retina to Light
- •Other Factors Contributing to Accelerated Accumulation of RPE Lipofuscin
- •A Hypothetical Scenario of Biogenesis of RPE Lipofuscin
- •Effects of Lipofuscin on RPE Function and Viability
- •Photoreactivity of RPE Lipofuscin
- •Toxicity of RPE Lipofuscin
- •Effects of Lipofuscin Components and Oxidative Stress in the RPE on Proinflammatory and Angiogenic Signaling
- •Approaches to Diminish Lipofuscin Accumulation or Lipofuscin-Induced Damage
- •Conclusions
- •References
- •A Nonspecific System Provides Nonphotic Information for the Biological Clock
- •Introduction
- •Nonphotic Information
- •Nonspecific Systems
- •Ascending Reticular-Activating System
- •Orexin/Hypocretin Projection
- •Intergeniculate Leaflet of the Thalamus
- •Anatomy
- •The Pharmacology of the IGL
- •Chronobiology
- •The Electrophysiology of the IGL
- •IGL as an Integrator of Photic and Nonphotic Information
- •Conclusions
- •References
- •The Circadian Clock: Physiology, Genes, and Disease
- •Introduction
- •Circadian Rhythms in Physiology and Behavior
- •Circadian Rhythms in Visual Function
- •Entrainment
- •Anatomy
- •The Suprachiasmatic Nucleus
- •Inputs to the SCN
- •Peripheral Oscillators
- •A Clock in the Eye
- •Oscillators Outside the Nervous System
- •Clock Genes
- •Human Implications
- •Summary
- •References
344 |
Sharpe and Stockman |
mean values Lmean, with the appropriate L(ser180) or L(ala180) spectral sensitivity values. These are provided in the Appendix of Stockman, Jägle, Pirzer, and Sharpe [47].
Directional Sensitivity
Variation of the point of entry of light entering the pupil (or the obliquity of the light rays reaching the retina) produces, besides the effective change of retinal illumination level (the Stiles-Crawford effect of the first kind [150]), a modification of hue (the Stiles-Craw- ford effect of the second kind [151]). There is a hue shift throughout the spectrum and an increase in saturation in the blue-green [152] spectrum, both of which will have minor (and difficult to calculate) consequences for photopic luminous efficiency under normal (free) viewing conditions. For scotopic luminous efficiency, the effects of the angle of incidence can be disregarded. This is because the directional sensitivity of the rod photoreceptors is very small in magnitude and virtually negligible in wavelength dependency [153].
Variations in the Contribution of Chromatic Channels
An age-related reduction of efficiency has been observed at long wavelengths for photopic luminous efficiency functions based on HBM [104]. The decline has been attributed to a reduced contribution of the chromatic pathway signals to brightness with age. Interestingly, some evidence suggests that the infant’s luminosity function is mediated by luminance (largely without a chromatic pathway contribution) rather than by brightness (with a chromatic pathway contribution) signals, although this is difficult to test [154]. To date, there remains no evidence that the infant visual system computes a brightness signal. Moreover, such young infants also fail to make chromatic discriminations. This may mean that the chrominance pathways develop later than the luminance pathway, which seems to be intact even in 8-week-old infant subjects (see [154]).
CONCLUSIONS
Spectral luminous efficiency is well defined for almost all conditions of scotopic (rod) vision by the CIE V′(λ) function [32]. For photopic vision, a luminous efficiency function that obeys additivity has been defined for neutral daylight conditions, V*(λ) [44]. It is valid for small central viewing fields of 2° or less in diameter. For large viewing fields, 10° ore more in diameter, the V10*(λ) may be preferred instead. Moreover, both V*(λ) and V10*(λ) are representative of techniques that obey Abney’s law of photometric additivity. For some conditions, for instance, those requiring side-by-side brightness matching, a function based on brightness photometry may be more appropriate (such as the CIE brightness-matching function). However, luminous efficiencies estimated by that function will not obey additivity. For mesopic vision, uncertainties about how the rod and cone signals interact to determine luminous efficiency and how their interaction changes with increasing luminance make any recommendations about which luminous efficiency function to use impracticable. The standard luminous efficiency functions generally apply to young observers for fixed viewing and measuring techniques. However, formulas are available for correcting them so that they more closely approximate the luminous efficiencies of any individual observer for other viewing conditions.
Luminous Efficiency Functions |
345 |
REFERENCES
1.Hecht S, Schlaer S, Pirenne MH. Energy, quanta and vision. J Opt Soc Am 1942;38:196–208.
2.Rodieck RW. The first steps in seeing. Sunderland, MA: Sinauer; 1998.
3.Schrödinger E. Über das Verhältnis der Vierfarben zur Dreifarbentheorie. Sitzungberichte Abt 2a, Mathematik, Astronomie, Physik, Meteorologie und Mechanik Akademie der Wissenschaften in Wien, Mathematisch-Naturwissenschaftliche Klasse 1925;134:471.
4.Luther R. Aus dem Gebiet der Farbreizmetrik. Z Tech Phys 1927;8:540–558.
5.Walls GL. A branched-pathway schema for the color-vision system and some of the evidence for it. Am J Ophthalmol 1955;39:8–23.
6.De Lange H. Research into the dynamic nature of the human fovea-cortex systems with intermittent and modulated light. II. Phase shift in brightness and delay in color perception. J Opt Soc Am 1958;48:784–789.
7.Guth SL, Alexander JV, Chumbly JI, Gillman CB, Patterson MM. Factors affecting luminance additivity at threshold. Vision Res 1968;8:913–928.
8.Smith VC, Pokorny J. Spectral sensitivity of the foveal cone photopigments between 400 and 500 nm. Vision Res 1975;15:161–171.
9.Boynton RM. Human color vision. New York: Holt, Rinehart and Winston; 1979.
10.Eisner A, MacLeod DIA. Blue sensitive cones do not contribute to luminance. J Opt Soc Am 1980;70:121–123.
11.Le Grand Y. Light, colour and vision. 2nd ed. London: Chapman and Hall; 1968.
12.Wyszecki G, Stiles WS. Color science: concepts and methods, quantitative data and formulae. 2nd ed. New York: Wiley; 1982.
13.Abney WdW. Festing ER. Colour photometry. Philos Trans R Soc Lond 1886;177:423–456.
14.Abney WdW. Researches in colour vision. London: Longmans, Green; 1913.
15.Dresler A. The non-additivity of heterochromatic brightness. Trans Illuminating Eng Soc 1953;18:141–65.
16.Boynton RM, Kaiser P. Vision: the additivity law made to work for heterochromatic photometry with bipartite fields. Science 1968;161:366–368.
17.Guth SL, Donley NV, Marrocco RT. On luminance additivity and related topics. Vision Res 1969;9:537–575.
18.Le Grand Y. Spectral luminosity. In: Jameson D, Hurvich LH, eds. Visual psychophysics, handbook of sensory physiology. Berlin: Springer-Verlag; 1972:413–433.
19.Wagner G, Boynton RM. Comparison of four methods of heterochromatic photometry. J Opt Soc Am 1972;62:1508–1515.
20.Pokorny J, Smith VC, Lutze M. Heterochromatic modulation photometry. J Opt Soc Am A 1989;6:1618–1623.
21.Lennie P, Pokorny J, Smith VC. Luminance. J Opt Soc Am A 1993;10:1283–1293.
22.Stockman A, Sharpe LT. Cone spectral sensitivities and color matching. In: Gegenfurtner K, Sharpe LT, eds. Color vision: from genes to perception. Cambridge, UK: Cambridge University Press; 1999:53–87.
23.Stockman A, Sharpe LT. Spectral sensitivities of the middleand long-wavelength sensitive cones derived from measurements in observers of known genotype. Vision Res 2000;40:1711–1737.
24.Eisner A, MacLeod DIA. Flicker photometric study of chromatic adaptation: selective suppression of cone inputs by colored backgrounds. J Opt Soc Am 1981;71:705–718.
25.Cavanagh P, MacLeod DIA, Anstis SM. Equiluminance: spatial and temporal factors and the contribution of blue-sensitive cones. J Opt Soc Am A 1987;4:1428–1438.
26.König A. Über den menschlichen Sehpurpur und seine Bedeutung fur das Sehen. Acad Wiss Sitzungsber 1894;30:577–598.
346 |
Sharpe and Stockman |
27.Willmer EN. Further observations on the properties of the central fovea in colour-blind and normal subjects. J Physiol (Lond) 1950;110:422–446.
28.Thomson LC, Wright WD. The colour sensitivity of the retina with the central fovea of man. J Physiol (Lond) 1947;105:316–331.
29.Williams DR, MacLeod DIA, Hayhoe MM. Foveal tritanopia. Vision Res 1981;19:1341–1356.
30.Wald G. Human vision and the spectrum. Science 1945;101:653–658.
31.Crawford BH. The scotopic visibility function. Proc Phys Soc Lond 1949;B62:321–334.
32.CIE. Commission Internationale de l’ Éclairage Proceedings; 1951. Vienna, Austria: CIE.
33.De Vries H. The luminosity curve of the eye as determined by measurements with the flicker photometer. Physica 1948;14:319–348.
34.Ikeda M, Urakubo M. Flicker HRTF as test of color vision. J Opt Soc Am 1968;58:27–31.
35.Marks LE, Bornstein MH. Spectral sensitivity by constant CFF: effect of chromatic adaptation. J Opt Soc Am 1973;63:220–226.
36.King-Smith PE, Webb JR. The use of photopic saturation in determining the fundamental spectral sensitivity curves. Vision Res 1974;14:421–429.
37.Eisner A. Comparison of flicker-photometric and flicker-threshold spectral sensitivities while the eye is adapted to colored backgrounds. J Opt Soc Am 1982;72:517–518.
38.Stromeyer CF III, Cole GR, Kronauer RE. Chromatic suppression of cone inputs to the luminance flicker mechanisms. Vision Res 1987;27:1113–1137.
39.Swanson WH. Chromatic adaptation alters spectral sensitivity at high temporal frequencies. J Opt Soc Am A 1993;10:1294–1303.
40.Stockman A, MacLeod DIA, Vivien JA. Isolation of the middleand long-wavelength sensitive cones in normal trichromats. J Opt Soc Am A 1993;10:2471–2490.
41.Stromeyer CF, III, Chaparro A, Tolias AS, Kronauer RE. Colour adaptation modifies the long-wave versus middle-wave cone weights and temporal phases in human luminance (but not red-green) mechanism. J Physiol (Lond) 1997;499:227–254.
42.CIE. Commission Internationale de l’Éclairage proceedings, 1924. Cambridge, UK: Cambridge University Press; 1926.
43.Gibson KS, Tyndall EPT. Visibility of radiant energy. Sci Pap Bureau Stand 1923;19:131–191.
44.Sharpe LT, Stockman A, Jagla W, Jägle H. A luminous efficiency function, V*(λ), for daylight adaptation. J Vis 2005;5:948–968.
45.Judd DB. Report of U.S. Secretariat Committee on Colorimetry and Artificial Daylight. In: Technical Committee No. 7 ed. Proceedings of the twelfth session of the CIE, Stockholm. Paris: Bureau Central de la CIE; 1951:1–60.
46.Vos JJ. Colorimetric and photometric properties of a 2-deg fundamental observer. Color Res Appl 1978;3:125–128.
47.Stockman A, Jägle H, Jagla W, Sharpe LT. The dependence of luminous efficiency on chromatic adaptation. J Vis. in press.
48.Ikeda M, Yaguchi H, Sagawa KJ. Brightness luminous-efficiency functions for 2 degrees and 10 degrees fields. J Opt Soc Am A 1982;72:1660–1665.
49.Ikeda M, Nakano Y. Spectral luminous-efficiency functions obtained by direct heterochromatic brightness matching for point sources and for 2 degrees and 10 degrees fields. J Opt Soc Am A 1986;3:2105–2108.
50.Stockman A, Sharpe LT. Into the twilight zone: the complexities of mesopic vision and luminous efficiency. Ophthal Physiol Opt 2006;26:225–239.
51.Aguilar M, Stiles WS. Saturation of the rod mechanism of the retina at high levels of stimulation. Opt Acta (Lond) 1954;1:59–64.
52.Adelson EA. Saturation and adaptation in the rod system. Vision Res 1982;22:1299–1312.
Luminous Efficiency Functions |
347 |
53.Bauer GM, Frumkes TE, Holstein GR. The influence of rod light and dark adaptation upon rod-cone interaction. J Physiol (Lond) 1983;337:121–135.
54.Bauer GM, Frumkes TE, Nygaard RW. The signal-to-noise characteristic of rod-cone interaction. J Physiol (Lond) 1983;337:101–119.
55.Frumkes TE, Sekuler MD, Reiss EH. Rod-cone interactions in human scotopic vision. Science 1972;175:913–914.
56.Frumkes TE, Sekuler MD, Barris MC, Reiss EH, Chalupa LM. Rod-cone interactions in human scotopic vision-I. Temporal analysis. Vision Res 1973;13:1269–1282.
57.Frumkes TE, Naarendorp F, Goldberg SH. The influence of cone adaptation upon rod mediated flicker. Vision Res 1986;26:1167–1176.
58.Alexander KR, Fishman GA. Rod-cone interaction in flicker perimetry. Br J Ophthalmol 1984;68:303–309.
59.Buck SL, Knight R, Hogden U. Why rod-cone interactions? J Opt Soc Am A 1985;2:P26.
60.Coletta NJ, Adams AJ. Rod-cone interaction in flicker detection. Vision Res 1984;24:1333–1340.
61.Frumkes TE, Temme LA. Rod-cone interactions in human scotopic vision-II. Cones influence rod increment thresholds. Vision Res 1977;17:673–679.
62.Goldberg SH, Frumkes TE, Nygaard RW. Inhibitory influence of unstimulated rods in the human retina: evidence provided by examining cone flicker. Science 1983;221:180–182.
63.Latch M, Lennie P. Rod-cone interaction in light adaptation. J Physiol (Lond) 1977;269:517–534.
64.Makous W, Boothe R. Cones block signals from rods. Vision Res 1974;14:285–294.
65.Shapiro AG. Cone-specific modulation of rod sensitivity in trichromatic observers. Invest Ophthalmol Vis Sci 2002;43:898–905.
66.Østerberg GA. Topography of the layer of rods and cones in the human retina. Acta Ophthalmol (Copenh) 1935;13(suppl 6):1–97.
67.Curcio CA, Sloan KR, Kalina RE, Hendrickson AE. Human photoreceptor topography. J Comp Neurol 1990;292:497–523.
68.König A. Die Abhängigkeit der Sehschärfe von der Beleuchtungsintensität. Acad Wiss Sitzungsber 1897;35:559–575.
69.Hecht S, Mintz EU. The visibility of single lines at various illuminations and the basis of visual resolution. J Gen Physiol 1939;22:593–612.
70.D’Zmura M, Lennie P. Shared pathways for rod and cone vision. Vision Res 1986;26: 1273–1280.
71.Hess RF, Nordby K, Pointer JS. Regional variation of contrast sensitivity across the retina of the achromat: sensitivity of human rod vision. J Physiol (Lond) 1987;388:101–119.
72.Hecht S, Shlaer S. Intermittent stimulation by light. V. The relation between intensity and critical frequency for different parts of the spectrum. J Gen Physiol 1936;19:965–977.
73.Arden GB, Weale RA. Variations of the latent period of vision. Proc R Soc B 1954;B142: 258–269.
74.Veringa F, Roelofs J. Electro-optical stimulation in the human retina. Nature 1966;211: 321–322.
75.MacLeod DIA. Rods cancel cones in flicker. Nature 1972;235:173–174.
76.Sharpe LT, Fach CC, Nordby K. Temporal summation in the achromat. Vision Res 1993;28:1263–1269.
77.van den Berg TJJP, Spekreijse H. Interaction between rod and cone signals studied with temporal sine-wave stimulation. J Opt Soc Am 1977;67:1210–1217.
78.Conner JD. The temporal properties of rod vision. J Physiol (Lond) 1982;332:139–155.
79.Sharpe LT, Stockman A, MacLeod DIA. Rod flicker perception: scotopic duality, phase lags and destructive interference. Vision Res 1989;29:1539–1559.
348 |
Sharpe and Stockman |
80.Stockman A, Sharpe LT, Zrenner E, Nordby K. Slow and fast pathways in the human rod visual system: ERG and psychophysics. J Opt Soc Am A 1991;8:1657–1665.
81.Stockman A, Sharpe LT, Rüther K, Nordby K. Two signals in the human rod visual system: a model based on electrophysiological data. Vis Neurosci 1995;12:951–970.
82.Walters HV, Wright WD. The spectral sensitivity of the fovea and parafovea in the Purkinje range. J Opt Soc Am 1943;45:507–514.
83.Kinney JAS. Spectral sensitivity of the eye to spectral radiation at scotopic, mesopic, and photopic intensity levels. J Opt Soc Am 1958;45:507–514.
84.Palmer DA. Standard observer for large-field photometry at any level. J Opt Soc Am 1968;58:1296–9.
85.Kokoschka S. Untersuchungen zur mesopischen Strahlungsbewertung. Die Farbe 1972;21: 39–112.
86.Nakano Y, Ikeda M. A model for brightness perception at mesopic levels. Kogaku (Japanese Journal of Optics) 1986;15:295–302.
87.Sagawa K, Takeichi K. Spectral luminous efficiency functions in the mesopic range. J Opt Soc Am A 1986;3:71–75.
88.Yaguchi H, Ikeda M. Mesopic luminous-efficiency functions for various adapting levels. J Opt Soc Am A 1984;1:120–123.
89.Viénot F, Chiron F. Brightness matching and flicker photometric data obtained over the full mesopic range. Vision Res 1992;32:533–540.
90.He Y, Bierman A, Rea MS. A system of mesopic photometry. Lighting Research and Technology 1998;30:175–181.
91.Kokoschka S, Bodmann HW. Ein konsistentes System zur photometrischen Strahlungsbewertung im gesamten Adaptationsbereich. In: Proceedings of the CIE 18th session, London, 1975. Vienna: Central Bureau of the CIE; 1975:217–225.
92.Trezona PW. A system of mesopic photometry. Color Res Appl 1991;16:202–216.
93.Ikeda M, Shimozono H. Mesopic luminous efficiency functions. J Opt Soc Am 1981;71: 280–284.
94.Bierman A, He Y, Rea MS. Visual reaction times: methods for measuring small differences. Lighting Res Technol 1998;30:169–174.
95.Sharpe LT, Stockman A, Jägle H, Nathans J. Opsin genes, cone photopigments, color vision and colorblindness. In: Gegenfurtner K, Sharpe LT, eds. Color vision: from genes to perception. Cambridge, UK: Cambridge University Press; 1999:3–51.
96.Stockman A, Sharpe LT, Merbs S, Nathans J. Spectral sensitivities of human cone visual pigments determined in vivo and in vitro. In: Palczewski K, ed. Vertebrate phototransduction and the visual cycle, Part B methods in enzymology, Vol. 316. New York: Academic Press; 2000:626–650.
97.Yaguchi H, Kawada A, Shioiri S, Miyake Y. Individual differences of the contribution of chromatic channels to brightness. J Opt Soc Am A 1993;10:1373–1379.
98.Stockman A, Sharpe LT, Fach CC. The spectral sensitivity of the human short-wavelength cones. Vision Res 1999;39:2901–2927.
99.van Norren D, Vos JJ. Spectral transmission of the human ocular media. Vision Res 1974;14: 1237–1244.
100.Sharpe LT, Stockman A, Jägle H, et al. Red, green, and red-green hybrid photopigments in the human retina: correlations between deduced protein sequences and psychophysicallymeasured spectral sensitivities. J Neurosci 1998;18:10053–10069.
101.Said FS, Weale RA. The variation with age of the spectral transmissivity of the living human crystalline lens. Gerontologia 1959;3:213–231.
Luminous Efficiency Functions |
349 |
102.Pokorny J, Smith VC, Lutze M. Aging of the human lens. Appl Optics 1988;26:1437–1440.
103.Kraft JM, Werner JS. Spectral efficiency across the life span: flicker photometry and brightness matching. J Opt Soc Am A 1994;11:1213–1221.
104.Sagawa KJ, Takahashi Y. Spectral luminous efficiency as a function of age. J Opt Soc Am A 2001;18:2659–2667.
105.Bieber ML, Volbrecht VJ, Werner JS. Spectral efficiency measured by heterochromatic flicker photometry is similar in human infants and adults. Vision Res 1995;35:1385–1392.
106.Xu J, Pokorny J, Smith VC. Optical density of the human lens. J Opt Soc Am A 1997;14: 953–960.
107.CIE. Fundamental chromaticity diagram with physiological axes—parts 1 and 2. Technical Report 17011. Vienna: Central Bureau of the Commission Internationale de l’ Éclairage; 2007.
108.Bone RA, Sparrock JMB. Comparison of macular pigment densities in the human eye. Vision Res 1971;11:1057–1064.
109.Pease PL, Adams AJ, Nuccio E. Optical density of human macular pigment. Vision Res 1987;27:705–710.
110.Bone RA, Landrum JT, Fernandez L, Tarsis SL. Analysis of the macular pigment by HPLC: retinal distribution and age study. Invest Ophthalmol Vis Sci 1988;29:843–849.
111.Moreland JD, Alexander EC. Effect of macular pigment on colour matching with field sizes in the 1° to 10° range. Documenta Ophthalmologica Proceedings Series 1997;59:363– 368.
112.Werner JS, Donnelly SK, Kliegl R. Ageing and human macular pigment density. Appended with translations from the work of Max Schultze and Ewald Hering. Vision Res 1987;27:257–268.
113.Chen S-F, Chang Y, Wu J-C. The spatial distribution of macular pigment in humans. Curr Eye Res 2001;23:422–434.
114.Ciulla TA, Curran-Celantano J, Cooper DA, et al. Macular pigment optical density in a midwestern sample. Ophthalmology 2001;108:730–737.
115.Broekmans WMR, Berendschot TTJM, Klöpping-Ketelaars IAA, et al. Macular pigment density in relation to serum and adipose tissue concentrations of Lutein and serum concentrations of Zeaxanthin. Am J Clin Nutr 2002;76:595–603.
116.Zagers NPA, van Norren D. Absorption of the eye lens and macular pigment derived from the reflectance of cone photoreceptors. J Opt Soc Am A 2004;21:2257–2268.
117.Terstiege H. Untersuchungen zum Persistenzund Koeffizientesatz. Die Farbe 1967;16:1–120.
118.Alpern M. Lack of uniformity in colour matching. J Physiol (Lond) 1979;288:85–105.
119.Miller SS. Psychophysical estimates of visual pigment densities in red-green dichromats. J Physiol (Lond) 1972;223:89–107.
120.King-Smith PE. The optical density of erythrolabe determined by retinal densitometry using the self-screening method. J Physiol (Lond) 1973;230:535–549.
121.King-Smith PE. The optical density of erythrolabe determined by a new method. J Physiol (Lond) 1973;230:551–560.
122.Smith VC, Pokorny J. Psychophysical estimates of optical density in human cones. Vision Res 1973;13:1199–1202.
123.Burns SA, Elsner AE. Color matching at high luminances: photopigment optical density and pupil entry. J Opt Soc Am A 1993;10:221–230.
124.Berendschot TTJM, van der Kraats J, van Norren D. Foveal cone mosaic and visual pigment density in dichromats. J Physiol (Lond) 1996;492:307–314.
125.Pokorny J, Smith VC, Starr SJ. Variability of color mixture data—II. The effect of viewing field size on the unit coordinates. Vision Res 1976;16:1095–1098.
350 |
Sharpe and Stockman |
126.Swanson WH, Fish GE. Age-related changes in the colour-match-area effect. Vision Res 1996;36:2079–2085.
127.De Vries H. Luminosity curves of trichromats. Nature 1946;157:736–737.
128.De Vries H. The heredity of the relative numbers of red and green receptors in the human eye. Genetica 1948;24:199–212.
129.Cicerone CM, Nerger JL. The relative numbers of long-wavelength-sensitive to middle- wavelength-sensitive cones in the human fovea centralis. Vision Res 1989;29:115–128.
130.Kremers J, Usui T, Scholl HPN, Sharpe LT. Cone signal contributions to ERGs in dichromats and trichromats. Invest Ophthalmol Vis Sci 1999;40:920–930.
131.Kremers J, Scholl HPN, Knau H, Berendschot TTJM, Sharpe LT. L/M-cone ratios in human trichromats assessed by psychophysics, electroretinography and retinal densitometry. J Opt Soc Am A 2000;17:517–526.
132.Carroll J, McMahon C, Neitz M, Neitz J. Flicker-photometric electroretinogram estimates of L:M cone photoreceptor ratio in men with photopigment spectra derived from genetics. J Opt Soc Am A 2000;17:499–509.
133.Albrecht J, Jägle H, Hood DC, Sharpe LT. The multifocal visual evoked potential (mfERG) and cone isolating stimuli: variation in L- and M-cone driven signals across the retina. J Vis 2002;2:543–558.
134.Carroll J, Neitz J, Neitz M. Estimates of L:M cone ratio from ERG flicker photometry and genetics. J Vis 2002;2:531–542.
135.Hofer H, Carroll J, Neitz J, Neitz M, Williams DR. Organization of the human trichromatic cone mosaic. J Neurosci 2005;25:9669–9679.
136.Crone RA. Spectral sensitivity in color-defective subjects and heterozygous carriers. Am J Ophthalmol 1959;48:231–238.
137.Rushton WAH, Baker HD. Red/green sensitivity in normal vision. Vision Res 1964;4:75–85.
138.Adam A. Foveal red-green ratios of normals, colorblinds and heterozygotes. Proc TelHashomer Hospital: Tel-Aviv 1969;8:2–6.
139.Vos JJ, Walraven PL. On the derivation of the foveal receptor primaries. Vision Res 1971;11:799–818.
140.Lutze M, Cox NJ, Smith VC, Pokorny J. Genetic studies of variation in Rayleigh and photometric matches in normal trichromats. Vision Res 1990;30:149–162.
141.Vimal RLP, Smith VC, Pokorny J, Shevell SK. Foveal cone thresholds. Vision Res 1989;29:61–78.
142.Dobkins KR, Thiele A, Albright TD. Comparisons of red-green equiluminance points in humans and macaques: evidence for different L:M cone ratios between species. J Opt Soc Am A 2000;17:545–556.
143.Gunther KL, Dobkins KR. Individual differences in chromatic (red/green) contrast sensitivity are constrained by the relative numbers of L- versus M-cones in the eye. Vision Res 2002;42:1367–1378.
144.Wesner MF, Pokorny J, Shevell SK, Smith VC. Foveal cone detection statistics in colornormals and dichromats. Vision Res 1991;31:1021–1037.
145.Brainard DH, Roorda A, Yamauchi Y, et al. Functional consequences of the relative numbers of L and M cones. J Opt Soc Am A 2000;17:607–614.
146.Sharpe LT, de Luca E, Hansen T, Jägle H, Gegenfurtner K. Advantages and disadvantages of human dichromacy. J Vis 2006;6:213–23.
147.Winderickx J, Battisti L, Hibiya Y, Motulsky AG, Deeb SS. Haplotype diversity in the human red and green opsin genes: evidence for frequent sequence exchange in exon 3. Hum Mol Genet 1993;2:1413–1421.
Luminous Efficiency Functions |
351 |
148.Neitz M, Neitz J. Molecular genetics and the biological basis of color vision. In: Backhaus WGK, Kliegl R, Werner JS, eds. Color vision: perspectives from different disciplines. Berlin: Walter de Gruyter; 1998:101–119.
149.Sharpe LT, Stockman A, Jägle H, Knau H, Nathans J. L, M and L-M hybrid cone photopigments in man: deriving λmax from flicker photometric spectral sensitivities,. Vision Res 1999;39:3513–3525.
150.Stiles WS. The directional sensitivity of the retina and the spectral sensitivity of the rods and cones. Proc R Soc B 1939;B127:64–105.
151.Stiles WS. The luminous efficiency of monochromatic rays entering the eye pupil at different points and a new colour effect. Proc R Soc B 1937;B123:90–118.
152.Enoch JM, Stiles WS. The colour change of monochromatic light with retinal angle of incidence. Opt Acta (Lond) 1961;8:329–358.
153.Van Loo JA, Jr, Enoch JM. The scotopic Stiles-Crawford effect. Vision Res 1975;15:1005–1009.
154.Teller DY, Pereverzeva M, Civan AL. Adult brightness vs. luminance as models of infant photometry: variability, biasability, and spectral characteristics for the two age groups favor the luminance model. JOV 2003;3:333–46.
155.Hood DC, Finkelstein MA. Sensitivity to light. In: Boff K, Kaufman L, Thomas J, eds. Handbook of perception and human performance. New York: Wiley; 1986:5-1–5-66.
