- •Contents
- •Acknowledgements
- •1 Introduction
- •1.1 Roots of the duplicity theory of vision: Ancient Greeks
- •1.2 Further development of the duplicity theory
- •Part I The development of the basic ideas of the duplicity theory from Newton to G. E. Müller
- •2 The Newton tradition
- •2.5 Conclusions
- •2.7 Maxwell: triplicity of colour vision proved
- •2.8 Helmholtz: the Young-Helmholtz colour theory
- •3 The Schultze tradition
- •3.1 The duplicity theory of Max Schultze
- •3.2 Evidence in favour of the theory
- •3.3 One or several types of cone?
- •3.5 Boll: discovery of rhodopsin as a visual photopigment
- •3.7 Phototransduction of rhodopsin
- •3.9 The duplicity theory of Parinaud
- •3.12 The duplicity theory of von Kries
- •1. Lights that match in day vision may differ in twilight vision: the Purkinje phenomenon.
- •2. Anatomical interpretation of the theory. Cones and Rods. Uniqueness of the fovea. Rhodopsin.
- •3. Isolation of twilight vision. Congenital, total colour-blindness. Nyctalopia. On comparative anatomy.
- •3.13 An attempt to unify the theories of Schultze and Young-Helmholtz
- •4 The Goethe tradition: the phenomenological approach
- •4.1 Phenomenological analysis may reveal underlying material processes
- •4.2 The colour theory of J. W. von Goethe
- •4.4 The colour theory of Ewald Hering
- •4.6 Contributions of Hering
- •5.1 The colour theory of Tschermak
- •5.2.2 Cones may inhibit regeneration of rhodopsin
- •5.2.3 Rods subserving chromatic colour vision
- •5.2.4 Three types of cones and five pairs of opponent processes
- •5.2.5 Activation of opponent processes by P1, P2 and P3
- •5.2.6 The P1 system
- •5.2.7 The P2 system
- •5.2.8 The P3 system
- •6 The duplicity theory of Polyak
- •6.1 Trichromacy of colour vision explained by three types of bipolar cell
- •6.2 Midget ganglion cells as synthesizers
- •6.3 The specific fibre-energy doctrine questioned
- •6.5 Common pathways of rods and cones
- •6.6 Explanations of acuity and sensitivity differences between rods and cones
- •6.7 The functional potentials of the synaptic arrangement
- •7.1 The electrical responses to light stimuli in single optic nerve fibres
- •7.2 The electrical responses in single optic nerve fibres of Limulus
- •7.3 The electrical responses in single optic nerve fibres of the frog
- •8 The duplicity theory of R. Granit
- •8.1 Supporting evidence for the duplicity theory from the ERG technique
- •8.2 The dominator-modulator theory
- •8.2.1. The trichromatic colour theory challenged
- •9.1 The duplicity theory of Willmer
- •9.1.1 Colour vision explained by two types of rod and one type of cone
- •9.3 Ivar Lie: interactions between rod and cone functions at mesopic intensity
- •9.3.1 Psychophysical experiments
- •9.3.2 The colour-mixing hypothesis
- •9.3.3 An alternative explanatory model
- •10 Status of the duplicity theory in the mid 1960s and its further development
- •Part III Chromatic rod vision: a historical account
- •11 Night vision may appear bluish
- •12 Mechanisms of chromatic rod vision in scotopic illumination
- •12.1 All principle hues may be observed in scotopic vision
- •12.2 Scotopic contrast colours are triggered by rod signals
- •12.3 Scotopic contrast colours depend on selective chromatic adaptation of cones
- •12.4 Scotopic hues explained
- •13 Rod-cone interactions in mesopic vision
- •13.1 Rod-cone interactions under mesopic conditions in a chromatically neutral state of adaptation
- •13.2 Rod-cone interactions under mesopic conditions in a chromatic state of adaptation
- •14 Further exploration of chromatic rod vision
- •14.1 Contribution of J. J. McCann and J. L. Benton
- •14.2 Contribution of P. W. Trezona
- •14.3 Contribution of C. F. Stromeyer III
- •14.4 Contribution of S. Buck and co-workers
- •14.5. Contribution of J. L. Nerger and co-workers
- •Part IV Theories of sensitivity regulation of the rod and cone systems: a historical account
- •15 Introduction
- •16 Early photochemical explanations
- •17 Contribution of S. Hecht
- •17.2 Supporting evidence obtained from invertebrates
- •17.3 Supporting evidence obtained from psychophysical experiments
- •18 Contribution of G. Wald: photochemical sensitivity regulation mechanisms of rods and cones
- •18.2 Serious challenges to the photochemical theory
- •18.3 The neural factor refuted
- •19 Relationship between amount of rhodopsin and sensitivity during dark adaptation
- •19.1 Results of Tansley
- •19.2 Results of Granit
- •19.5 A logarithmic relationship between sensitivity and amount of bleached photopigment
- •19.7 Contribution of W.A.H. Rushton: relationship between sensitivity and amount of bleached rhodopsin in humans
- •20 Post-receptor sensitivity regulation mechanisms
- •20.1 Psychophysical evidence
- •20.2 Anatomical and electrophysiological evidence
- •21.1 Each receptor type has a separate and independent adaptation pool
- •21.2 Are light and dark adaptation really equivalent?
- •21.3 A decisive experiment
- •21.4 The adaptation mechanisms explored by the after-flash technique
- •22 Contribution of H. B. Barlow
- •22.1 Dark and Light adaptation based on similar mechanisms
- •22.2 Both noise and neural mechanisms involved
- •22.3 Evidence in support of the noise theory
- •22.4 Opposing evidence
- •22.5 Sensitivity difference between rods and cones explained
- •23 Rushton and Barlow compared
- •24.1 Contribution of T.D. Lamb
- •24.2 The search for a new formula
- •24.3 Differences between rod and cone dark adaptation
- •24.4 Light and dark adaptation are not equivalent
- •24.5 Allosteric regulation of dark adaptation
- •24.6 A search for the allosteric adaptation mechanisms
- •25 Several mechanisms involved in sensitivity regulation
- •26 Sensitivity regulation due to rod-cone interaction
- •27 Modern conceptions of sensitivity regulation
- •Part V Factors that triggered the paradigm shifts in the development of the duplicity theory
- •References
- •Index
Index
absorption spectrum 144 achromatic sensation 28, 30, 83
acuity performance 25, 30, 31, 69, 70, 71 adaptation pool 163
additive opponent hue 118 ‘after-blueness’ 126
AGC pool 157, 160, 162, 174 alcohol dehydrogenase 143 alcohol group of vitamin A 143
algebraic treatment of the colour-mixture data 18
‘all-or-none’ law 81 allosteric proteins 181 all-trans retinal 140, 143 amacrine cells 65, 70, 193
amount of bleached rhodopsin 153, 178, 179, 180
‘Anagenese’ 29
analytical histological methods 62 Ancient Greeks 2, 3, 205 anomalous trichromats 87 apoenzyme 143
assimilation processes 48, 49 ‘Ausgangsmaterial’ 54, 59
binary colour mixtures 47 binocular colour mixture 56 binocular matching technique 187 bioelectrical output 71
bleaching-regeneration cycle 140, 142 Bunsen-Roscoe law 73, 167
b-wave 80, 148, 151, 152
Cajal’s idea 67
carbon-to-carbon double bond 141 carbonyl group 143 carotenoid-protein 140, 142 centripetal impulses 71
chicken retina 141, 142 chimpanzees 62 chromophore 143 classical logic 199
coefficient law 187 coenzymes 143
colour circle 12, 13, 44, 45, 197 common rod and cone pathways 67 comparative histological studies 22 compartment theory 149, 150, 151, 153 concept of light 10
conformational alteration 182 corpuscular theory of light 10 ‘Crawford-Westheimer effect’ 189 crystalline lens 205
cyclase activity 183, 184, 185
cyclic guanosine monophosphate 183 cytoplasmic face 184
‘dark light’ 171, 173, 176
‘das allgemeine psychophysische Grundgesetz’ 47
‘day rods’ 88, 90, 105 deductive reasoning 199 deutranope 90, 109 dissimilation processes 48 diurnal 23, 24, 132, 195, 203
dominator and modulator curves 82 dominator curve 82 dominator-modulator theory 83 double reciprocal plots 180
double slit experiment 10 Dowling-Rushton equation 166, 176,
177
electrical activity of the fibre 72 electrodes 78 electroretinogram 78
empty space 10 epoch-making discoveries 26 E-retina 79, 80 ERG-technique 78, 179 ether-like substance 2 evolutionary development 89 excitatory cascade 185 extreme periphery 90
222 index
falsifiable theories 199 falsification tests 199, 200 ‘fast’ rod influence 128 flickering light 96, 98 four basic elements 2 frequency of discharge 74
fresh-water vertebrates 141 frog retina 74, 90
functional organization of receptive fields 75
galvanometer 78 ‘Goethe tradition’ 5 Golgi method 62, 69, 70 guanylate cyclase 183
‘héméralopie’ 31, 134
Hering’s colour theory 49, 117, 119 highest acuity 65
histological evidence 190 hydrogen atoms 143
hyperpolarizing change of the plasma membrane potential 185
identical subunits 181, 184 ‘incommensurable’ 201 interference pattern 10
intramolecular model of dark adaptation 182 intricate synaptic pattern 70
intrinsic noise 169 invertebrates 26 iodopsin 141, 142 I-retina 79, 80
isomerize the chromophore 142
‘lantern’ theory 2
lateral geniculate nucleus 117 laws of refraction 195
L-cone pathways 129 ligand concentration 181
light-sensitive ion channel 184 ‘likeness’ principle 2
Limulus 72, 73, 74
macaque monkey 117 Mach’s maxim 41 marine-fish species 141
McCollough colour after-effect 127, 189 M-cone pathways 129
mechanisms of phototransduction 185 messenger particles 2
meta II 143, 166
microelectrodes 41, 61 microelectrode technique 78, 81 midget ganglion cells 65
mixed rod-cone retina 79, 81, 82, 89 modern colorimetry 12
mop, brush, flat, and midget bipolar 64 multicoloured image 124, 125 multiple retinal gain controls 192
negatively cooperative 180 ‘Neogenese’ 29
nerve webs 70
Newton’s law of gravity 18 ‘Newton tradition’ 4, 7 night-blind 31
night vision 23, 30, 32, 106, 132, 134, 193 nocturnal 36, 96, 132, 195, 203, 244 ‘normal’ scientific period 200, 204 nucleotide sequence analysis of cloned
DNA 185
occipital cortex 145
oligomeric protein complex 184 opponency 50
opsin 142, 143 optical sensitizer 54 optic nerve 11 ‘optimum colour’ 125
‘optochemische Hypothese’ 25 oscillograph 72
oxidized 143
paradigm shift 20, 22, 26, 50, 196 PDE hydrolysis 185
phenomenological analysis 45, 46, 47, 49, 52, 197
phosphodiesterase 183 phosphorylation enzymes 193 photoelectric effect 10 ‘photon-like’ events 169 pigment epithelium 143, 144 Planck’s quantum principle 10 plasma membrane 184 porphyropsin 141, 142
positively cooperative protein 184 pre-scientific period 200, 201 principle of gravitation 12, 14, 197 Principle of Univariance 73, 108, 115 prismatic solar spectrum 195 protanope 18, 90
pupil size 186
Purkinje afterimage 38, 92 Purkinje phenomenon 31, 33, 53, 93 Purkinje shift 36, 79, 82, 96, 142
‘quanta’ of light 10
rapid voltage fluctuations 72 receptive field 74, 76, 157 ‘red-green’ see-substance 48 retinoids 193
rod-cone facilitation 121
rod monochromat 33, 54, 55, 177 roots of the duplicity theory 3
‘Schultze trawdition’ 4 scientific revolution 201 S-cone pathways 129
scotopic contrast colours 113, 114, 116, 118, 119
sensorium 11, 34
simultaneous and successive contrast 49 simultaneous and successive induction 49 single binding site for cGMP 185
‘slow’ rod influence 128
specific fibre-energy doctrine 34, 84 specific-hue threshold 101, 102, 103, 104,
111, 113, 120, 121, 122 specific nerve-energy doctrine 20 spectrally opponent cells 117, 121 spectral modulator curves 82
spectro-photometrical measurements 111 Spinoza’s principle of psychophysical
parallelism 41
index 223
statistical fluctuation 169, 174 Stiles-Crawford effect 125 succession of falsifications 200 succession of traditions 200
tetramer 181, 184
three standard lights 17, 18, 19
time constant of rhodopsin regeneration 178 transducin 182, 183
transient tritanopia 188 transmembrane ion flux 185 transverse pulses 9
trichromatic colour theory of Young 16, 20 two-colour threshold technique 93, 161 two-dimensional colour space 19
ultimate truth 199, 201, 206 unexpected observations 205 universal colour theory 14 ‘Urfarben’ 59, 119
visual yellow 29, 32, 33, 34, 88 vitamin A 140, 143, 144
‘water’ element 2
‘white-black’ see-substance 48, 49 white-grey-black colour series 47
Whiteness 8
‘yellow-blue’ see-substance 48, 49 yellow-blue substrate 34
Young- Maxwell-Helmholtz colour theory 206