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1. Direct measurements of the minimum energy required for threshold vision under optimal physiological conditions yield values between 2.1 and 5.7 x 10–10 ergs at the cornea, which correspond to between 54 and 148 quanta of blue-green light.
2. These values are at the cornea. To yield physiologically significant data they must be corrected for corneal reflection, which is 4 per cent; for ocular media absorption, which is almost precisely 50 per cent; and for retinal transmission, which is at least 80 per cent. Retinal transmission is derived from previous direct measurements and from new comparisons between the percentage absorption spectrum of visual purple with the dim-vision luminosity function. With these three corrections, the range of 54 to 148 quanta at the cornea becomes as an upper limit 5 to 14 quanta actually absorbed by the retinal rods.
3. This small number of quanta, in comparison with the large number of rods (500) involved, precludes any significant two quantum absorptions per rod, and means that in order to produce a visual effect, one quantum must be absorbed by each of 5 to 14 rods in the retina.
4. Because this number of individual events is so small, it may be derived from an independent statistical study of the relation between the intensity of a light flash and the frequency with which it is seen. Such experiments give values of 5 to 8 for the number of critical events involved at the threshold of vision. Biological variation does not alter these numbers essentially, and the agreement between the values measured directly and those derived from statistical considerations is therefore significant.
5. The results clarify the nature of the fluctuations shown by an organism in response to a stimulus. The general assumption has been that the stimulus is constant and the organism variable. The present considerations show, however, that at the threshold it is the stimulus which is variable, and that the properties of its variation determine the fluctuations found between response and stimulus.
Submitted on March 30, 1942
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|
|
|
|
|
|
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|
|
|
|
 |
|
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|
|
|
|
|
|
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|
|
|
|
|
|
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|
|
|
|
 |
|
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|
|
|
|
 |
|
 M. Han, J. Lou, K. Nakanishi, T. P. Sakmar, and S. O. Smith Partial Agonist Activity of 11-cis-Retinal in Rhodopsin Mutants J. Biol. Chem., September 12, 1997; 272(37): 23081 - 23085. [Abstract] [Full Text] [PDF]
|
|
|
|
|
|
D. O'Brien The chemistry of vision Science, December 3, 1982; 218(4576): 961 - 966. [Abstract] [PDF]
|
|
|
|
|
|
A Fein and D. Corson Both photons and fluoride ions excite limulus ventral photoreceptors Science, April 6, 1979; 204(4388): 77 - 79. [Abstract] [PDF]
|
|
|
|
 |
|
 D. A. Baylor and R. Fettiplace Transmission of Signals from Photoreceptors to Ganglion Cells in the Eye of the Turtle Cold Spring Harb Symp Quant Biol, January 1, 1976; 40(0): 529 - 536. [Abstract] [PDF]
|
|
|
|
|
|
G. L. Fain, G. H. Gold, and J. E. Dowling Receptor Coupling in the Toad Retina Cold Spring Harb Symp Quant Biol, January 1, 1976; 40(0): 547 - 561. [Abstract] [PDF]
|
|
|
|
|
|
G. Fain Quantum sensitivity of rods in the toad retina Science, March 7, 1975; 187(4179): 838 - 841. [Abstract] [PDF]
|
|
|
|
|
|
W. H. Miller, R. E. Gorman, and M. W. Bitensky Cyclic Adenosine Monophosphate: Function in Photoreceptors Science, October 15, 1971; 174(4006): 295 - 297. [Abstract] [PDF]
|
|
|
|
|
|
T. F. Budinger, H. Bichsel, and C. A. Tobias Visual Phenomena Noted by Human Subjects on Exposure to Neutrons of Energies Less than 25 Million Electron Volts Science, May 21, 1971; 172(3985): 868 - 870. [Abstract] [PDF]
|
|
|
|
|
|
G. Wald Molecular Basis of Visual Excitation Science, October 11, 1968; 162(3850): 230 - 239. [PDF]
|
|
|
|
|
|
R. A. Cone Early Receptor Potential: Photoreversible Charge Displacement in Rhodopsin Science, March 3, 1967; 155(3766): 1128 - 1131. [Abstract] [PDF]
|
|
|
|
|
|
G. Wald Visual Excitation and Blood Clotting Science, November 19, 1965; 150(3699): 1028 - 1030. [Abstract] [PDF]
|
|
|
|
|
|
W. A. Hagins Electrical Signs of Information Flow in Photoreceptors Cold Spring Harb Symp Quant Biol, January 1, 1965; 30(0): 403 - 418. [Abstract] [PDF]
|
|
|
|
|
|
W. E. Reichardt Quantum Sensitivity of Light Receptors in the Compound Eye of the Fly Musca Cold Spring Harb Symp Quant Biol, January 1, 1965; 30(0): 505 - 515. [Abstract] [PDF]
|
|
|
|
|
|
J. Scholes Discontinuity of the Excitation Process in Locust Visual Cells Cold Spring Harb Symp Quant Biol, January 1, 1965; 30(0): 517 - 527. [Abstract] [PDF]
|
|
|
|
|
|
C. H. Graham and Y. Hsia Color Defect and Color Theory: Studies of normal and color-blind persons, including a subject color-blind in one eye but not in the other. Science, March 28, 1958; 127(3300): 675 - 682. [Abstract] [PDF]
|
|
|
|
|
|
J. C. PESKIN Concentration of Visual Purple in a Retinal Rod of Rana pipiens Science, January 11, 1957; 125(3237): 68 - 69. [PDF]
|
|
|
|
|
|
G. Wald On the Mechanism of the Visual Threshold and Visual Adaptation Science, June 25, 1954; 119(3104): 887 - 892. [PDF]
|
|
|
|
|
|
G. WALD HUMAN VISION AND THE SPECTRUM Science, June 29, 1945; 101(2635): 653 - 658. [PDF]
|
|
|
|
