- •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
50development of the basic ideas of the duplicity theory
an achromatic light in the same area, the achromatic light appears greenish. In this case, the green colour observed was explained by Hering in a straightforward manner by the assumption that the sensitivity of the ‘red’ dissimilation process compared to that of the ‘green’ assimilation process was reduced in the stimulated area during the pre-stimulation period.
4.6 Contributions of Hering
It can be seen that Hering agreed with Goethe that opponency is a fundamental characteristic of colour phenomena, but in addition he also postulated an opponency of underlying material events. This clearly opposes the suggestion of the Young-Helmholtz colour theory (1896) that the nervous pathways, from receptor to brain, are like isolated telegraph wires with no interaction between different neural processes.
With hindsight, we can see that Hering pointed in the right direction. Indeed, his ingenious ideas deserve to be ranked as the fifth major paradigm shift in vision research. Yet, his phenomenological approach has two serious weaknesses. Firstly, the phenomenological report gives conclusive information only on the relative values of the underlying material processes. In the case of the successive contrast experiment, for instance, where the red pre-stimulation produced a green after-effect upon test stimulation, Hering could reasonably presume that the sensitivity for dissimilation relative to that of assimilation was reduced in the red-green see-substance. But even if this presumption is accepted, he was not in a position to decide whether the sensitivity for assimilation in the test area increases or remains constant. The sensitivity might even decrease, although less so than for dissimilation (Hering, 1878, p. 86).
Secondly, presupposing that the psychophysical maxim of Mach (1865) is valid, one may draw legitimate conclusions about material processes directly underlying consciousness, but not about visual processes prior to the final material process. This limitation of the approach is clearly revealed when the conditions are arranged so
the goethe tradition: the phenomenological approach 51
that test stimulation of rods and cones initiates the same achromatic colour. Since the subject cannot discriminate between the achromatic colours obtained under photopic and scotopic conditions, it follows from the maxim of Mach that the underlying material events are the same. Hence, the researcher may be misled into neglecting the important retinal differences involved.
Of course, the finding that the rod and cone receptor systems may give rise to the same achromatic colour sensation was also a major challenge for the Young-Helmholtz trichromatic colour theory. Here, the white sensation was assumed to result from an equal stimulation of the three primary cone receptor systems, while the rod system did not have any significant role to play (see Helmholtz, 1867; von Kries, 1911).
These serious shortcomings of the Hering and Young-Helmholtz colour theories pointed to the need for a more comprehensive theory of colour vision where all the three major traditions were incorporated (i.e. the Newton, Schultze and Goethe traditions). This lead was followed by several research workers in the early 1900s. The most detailed and comprehensive theories were provided by Armin Tschermak (1902, 1929) and George Elias Müller (1896, 1897, 1923, 1930), both rooted in the Goethe tradition.
5 The colour theories of Armin
Tschermak and George Elias Müller
5.1 The colour theory of Tschermak
In developing his own colour theory, Tschermak (1902, 1929) made a critical evaluation of the colour theories of Young-Helmholtz, Schultze and Hering. With regard to Young-Helmholtz’s colour theory, he was severely critical. Thus, he asserted that the basic assumption of three independent, primary colour-related processes postulated by the trichromatic theory could not be reconciled with the phenomenological analysis of colour sensation that revealed six qualitatively different unitary sensations: red, yellow, green, blue, white and black. It would, for example, be impossible to give an adequate explanation of the uncompounded yellow-related material process by greenand red-related processes, or the uncompounded white-related process by red-, greenand violet-related processes. Also, in opposition to the trichromatic colour theory, experiments on colour mixture, colour induction and colour contrast clearly revealed opponent interaction processes going on in the visual system.
Finally, in accord with von Kries (1911), Tschermak (1902, 1929) pointed out that the basic assumption of the trichromatic colour theory, that white sensation was generated when the three different types of cone receptors were activated to about the same degree, was seriously challenged by the fact that colourless sensation could also be observed in scotopic vision where only rod receptors were known to function.
With regard to Schultze’s duplicity theory, on the other hand, he found the evidence strongly in favour of its basic assumptions that cones functioned in day vision giving rise to both achromatic and chromatic sensations, and that rods functioned in night vision, giving
the colour theories of a. tschermak and g. e. müller 53
rise to achromatic sensation only. He also accepted the basic assumption of Hering’s opponent colour theory that colour vision rested upon independent, opponent ‘red-green’, ‘yellow-blue’ and ‘whiteblack’ material processes. On the basis of his evaluation, he came to the conclusion that there were five different kinds of cone receptor systems in the retina: an achromatic-, a red-, a yellow-, a greenand a blue-related system, in addition to the achromatic rod system.
Yet, his most original theoretical contribution was his suggestion that rhodopsin was situated not only in rods but also in small amounts in cones (Tschermak, 1929, p. 576). He found supporting evidence in the fact that several research workers had obtained the Purkinje phenomenon at the central fovea (Tschermak, 1902, p. 720). That some researchers (see e.g. von Kries, 1929) were unable to repeat this observation, he explained by the suggestion that the Purkinje phenomenon at the fovea was less pronounced than that obtained extrafoveally.
The suggestion of Tschermak that cones also contained rhodopsin raised the important question of how stimulation of rhodopsin might influence chromatic colour vision – a question discussed at length by G. E. Müller.
5.2 The duplicity theory of G. E. Müller
Along with von Kries, G. E. Müller was generally considered to be the leading authority in vision research in the 1920s. His colour theory was much more detailed and comprehensive than that of Tschermak (see G. E. Müller, 1896, 1897, 1923, 1930). It incorporated the basic knowledge of rod and cone functions accumulated up to 1930 and may be seen to represent the end of the first phase in the development of the duplicity theory, integrating evidence from the Newton, Schultze and Goethe traditions.
His theory, though, is highly speculative. Thus, in sharp contrast to the cautious theory construction of von Kries (1911), it gives detailed descriptions of colour processes in the visual pathway based on an almost complete lack of factual knowledge of the underlying neurophysiological processes. With hindsight, one may ask whether
