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PHYSICAL BASIS FOR VISUAL PERCEPTION AND COLOR        23


                                                     505
                                       1.0                       Day vision:
                                                           555    (cones)
                                            Night vision:
                                              (rods)
                                      Relative response  0.5










                                       0.0
                                             400      500      600      700
                                                      Wavelength (nm)

                       Figure 2-6
                       The spectral response of the eye in night and day vision. The two curves have been
                       normalized to their peak sensitivity, which is designated 1.0; however, night (rod) vision is 40
                       times more sensitive than day (cone) vision. Rhodopsin contained in rod cells and color
                       receptor pigments in cone cells have action spectra with different maxima and spectral
                       ranges.

                          The shape and distribution of the light-sensitive rod and cone cells in the retina are
                       adapted for maximum sensitivity and resolution in accordance with the physical param-
                       eters of light and the optics of the eye-lens system. Namely, the outer segments of cone
                       cells, the cells responsible for color perception in the fovea, are packed together in the
                       plane of the retina with an intercellular spacing of 1.0–1.5  m, about one-half the radius
                       of the minimum spot diameter (3  m) of a focused point of light on the retina. The small
                       1.5  m cone cell diameter allows the eye to resolve structural details down to the theo-
                       retical limit calculated for the eye-lens system. For an object held 25 cm in front of the
                       eye, this corresponds to spacings of  0.1 mm. It appears nature has allowed the light
                       receptor cells to utilize the physics of light and the principles of lens optics as efficiently
                       as possible!
                          Rod cell photoreceptors comprise 95% of the photoreceptors in the retina and are
                       active in dim light, but provide no color sense. Examine Figure 2-1 showing the struc-
                       ture of the eye and Figure 2-7 showing the distribution of rod cells in the retina. Rods
                       contain the light-sensitive protein, rhodopsin, not the photovisual pigments required for
                       color vision, and the dim light vision they provide is called scotopic vision. Rhodopsin,
                       a photosensitive protein, is conjugated to a chromophore, 11-cis-retinal, a carotenoid
                       that photoisomerizes from a cis to trans state upon stimulation and is responsible for
                       electrical activity of the rod cell membranes. The peak sensitivity of the rod photore-
                       ceptor cells (510 nm) is in the blue-green region of the visual spectrum. Rod cell vision
                       is approximately 40 times more sensitive to stimulation by light than the cone cell
                       receptors that mediate color vision. Bright light rapidly bleaches rhodopsin, causing
                       temporary blindness in dim lighting conditions, but rhodopsin isomerizes gradually
                       over a 20–30 min period, after which rod receptor function is largely restored. Full
                       recovery may require several hours or even days—ask any visual astronomer or micro-
                       scopist! To avoid photobleaching your rhodopsin pigments and to maintain high visual
                       sensitivity for dim specimens (common with polarized light or fluorescence optics), you
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