Dark adaptation of the long-wave cones at different eccentricities

Using a Wright colorimeter the ordinary long-term, long-wave cone dark-adaptation curve was measured at 0, 2, 4, 7, 17, 25, 40 and 49 degrees nasally in the visual field. In opposition to previous findings, the results show that the dark-adaptation function of the long-wave cones changes markedly when the test field is moved outward from the rod-free fovea. It is suggested that the kinetics of the long-wave cone photopigment change with eccentricity. Also, at variance with previous findings, the present curves at all eccentricities may reasonably well be interpreted as consisting of three different sections; a first section where the threshold decreases rapidly, followed by a major, approximately linear section and a terminating section that converges asymptotically towards the final level of sensitivity. This finding suggests that the dark-adaptation process of the cone system, under the given experimental conditions, is based on three somewhat different processes.     

Dark-adaptation of the human rod system: A new hypothesis

Following a long, full bleach, rod dark-adaptation curves from two normal trichromats were obtained with test fields of various size, exposure time and retinal eccentricity. The results show that there is a substantial region of threshold recovery with an approximately constant, linear slope of about 0.27 log per minute of dark-adaptation, which is independent of the test variables. It is suggested that the increase in sensitivity during this constant, linear slope is completely determined by changes in the concentration of bleached rhodopsin. The relationship between change of relative threshold (T) and fraction of bleached rhodopsin ( B ) is given by T=B ^3,7. This exponential law is well described by the displacement of the equilibrium between the active and inactive states of an allosteric enzyme built as a tetrarner.     

Mechanisms of long-term dark adaptation

It has previously been suggested that long-term dark adaptation is controlled by bleaching signals that regulate the activity of an allosteric, positively cooperative protein (Stabell et al., 1986a, b). Recent biochemical evidence strongly supports this assumption, indicating that the primary regulator of the light-sensitive channels in the plasma membrane of the outer segments of the photoreceptors is a homo-oligomeric, allosteric, positively cooperative protein. In this report, we discuss the possibility that signals from bleached photopigments may control the dark-adaptation process through the allosteric protein of the plasma membrane. It is suggested that the concentrations of the bleached photopigment and of the allosteric effector are reciprocal quantities.     

RODS AS COLOR RECEPTORS IN PHOTOPIC VISION

Comparing a foveal and an extra-foveal field during dark adaptation, transition from chromatic to achromatic vision at intensity levels above the cone plateau started around the break of the dark adaptation curve. Pre-stimulating the two fields in a dark-adapted state with deep red, and test-stimulating when returning sensitivity had reached absolute threshold of the dark-adapted eye, with green filters at intensities above the specific threshold, the fields matched as to hue and saturation. It appears that rod and cone activities are integrated and function as a synchronized unit during the initial recovery phase of dark adaptation.     

Limulus psychophysics: dark adaptation in the ventral eye.

Sudden illumination applied to Limulus produces an unconditioned downward tail movement which is under stimulus control and can be used to measure psychophysical thresholds. The method of constant stimuli was used to measure the behavioral dark-adaptation function mediated by the ventral eye of Limulus. The resulting function has two phases, each of which is rectified when log threshold is made a function of long time in the dark. Under the conditions of the present experiment, the transition between the two phases occurred at 6 min. This psychophysical dark-adaptation function has the same form as an electrophysiologic dark-adaptation function obtained by Fein and DeVoe (using the ventral eye receptor potential as the response). In more complex visual systems such two-phase, dark-adaptation functions would usually be interpreted in terms of screening pigment movements or changes in the neural contributions of different receptor classes, but neither interpretation is appropriate for the ventral eye of Limulus.     

Long-term rod dark adaptation in man. Threshold measurements, rhodopsin regeneration and allosteric sensitivity regulation. An evaluation

Recent evidence strongly suggests that the relationship between threshold elevation ( T ) and fraction of bleached rhodopsin ( B ), obtained during a major, middle period of long-term rod dark adaptation in man, is well described by a power function, i. e., T = k · Bⁿ , where k is a multiplicative constant and n is the exponent. Due primarily to the low reliability of measurements of rhodopsin regeneration, however, the exponent n of the power function cannot, at present, be given an exact value. Available information indicates that the value of the exponent ranges between 2.4 and 4. Implications of this uncertainty are discussed within the framework of the allosteric, tetrameric model of rod dark adaptation. It is concluded that this model in its simplest form may only offer a first approximation of the real system implicated in the process.     

FACILITATION OF CHROMATIC CONE ACTIVITY BY ROD ACTIVITY

STABELL, U. & STABELL, B. Facilitation of chromatic cone activity by rod activity. II. Variation of chromatic-related cone activity. Scand. J. Psychol. , 1971, 12, 168–174.–At the cone-rod break of the dark adaptation curve, the specific threshold was found to drop to lower intensity levels, while the threshold curves of fovea proceeded in one step only, confirming the suggestion that rods may facilitate chromatic-related cone activity.     

Stereoscopic acuity for photometrically matched background wavelengths at scotopic and photopic levels

Two Ss made equidistance settings in a two-rod apparatus using white and four chromatic (red, yellow, Feen, and blue) backgrounds, photometrically matched at each of eight or nine teat levels within a total retinal-illuminance range of 4 log units. The binocular depth setting were analyzed in terms of the angular magnitude of both the Yariable error, η_AD, and the constant error, η_ΔR When η_AD is plotted as a function of retinal illuminance, the curves for each of the four chromatic IIIId white backlfound coditions show that at low retinal illuminances, η_AD, is initially large, and with increasing background illumination, η_AD progressively decreases to approach a final low asymptotic value. As predicted by the duplicity theory of vision, each experimental curve shows a dircontinulty at about ?1.0 lot td (corresponding to a background luminance value of about 0.0069 fL with the 2.5-mm artificial pupil used) reflecting the transition from rod to cone functioning. The curves representing the different wavelengths essentially overlap throughout the total illumination range, indicating that, for both rod and cone vision, wavelength has no differential effect on the variability of depth settings. The data for η_ΔR are less regular than those for η_AD and the rod-cone discontinuities appear less pronounced. The data are analyzed in terms of the relative contribution of the rod and cone mechanisms to performance level.     

Comparison of scotopic sensitivity in diurnal (Anas platyrhynchos) and crepuscular (Dendrocygna autumnalis) ducks.

Scotopic visual adaptation curves were obtained from 4 mallard ducks. A curve of best fit was used to compare the mallards' mean adaptation curve to the curve previously reported for the black-bellied tree duck, a crepuscular species. The curves did not differ significantly in either their slopes or base levels (thresholds). The mallards curve had a rod-cone "break" at approximately 25 min. This break is evident in the scotopic curve for pigeons, but is absent from the black-bellied tree ducks' curve. Examination of retinal tissues indicated that the black-bellied tree ducks had significantly more rods and cones, and a larger rod:cone ratio than the mallards. The mallards' scotopic visual threshold is exceeded by the natural illumination present under several nocturnal conditions.     

Rod involvement in peripheral color processing

Abstract.— It has previously been suggested that rods act as blue receptors in peripheral color vision. Two experiments examining this issue were conducted. Experiment 1 investigated the perceived hue of a test light presented at a luminance level above chromatic threshold. At 8° in the periphery, the 500 nm test light was perceived as more blue under conditions of dark adaptation than after light adaptation. Similar differences were not found for foveal presentation. The increased blue in the periphery after dark adaptation was attributed to a rod contribution. In Experiment 2 an attempt was made to mix a rod contribution obtained with a 470 nm light below chromatic threshold, with a cone color obtained from a 670 nm light presented above chromatic threshold. No evidence was obtained to support the idea that a blue produced by rods stimulated below chromatic threshold could mix with a red produced by cones stimulated above chromatic threshold. The results are discussed in terms of a rod contribution to hue which is dependent on the luminance level of short wavelength stimulation.     

TRANSITION FROM ROD TO CONE VISION:III. Successive contrast anew

S tabell , U. & S tabell , B. Transition from rod to cone vision. III. Successive contrast anew. Scand. J. Psychol ., 1969, 10 , 140–144—The relation between the specific threshold level and the upper limit of the scotopic contrast color was investigated. The achromatic interval between the scotopic and the photopic component increased when time in darkness increased, and when pre-stimulation was reduced as regards intensity, duration, or cone/rod ratio. The results are interpreted on the basis of the opponent theory of color vision.     

Brightness interactions between rods and cones

Two parafoveal test targets with different spectral compositions were matched in brightness to a fixed-luminance foveal reference target under scotopic adaptation conditions. The idea of the experiment was to find a reference luminance for which one of the matching test targets stimulated only rods while the other stimulated both rods and cones. If brightness was proportional to the linear sum of rod and cone responses, then the luminance of the matching rod+cone target would be predictably closer to rod threshold than would that of the rod target. The results were complicated by evidence that rod responses to the test targets selectively enhanced weak chromatic signals. Nevertheless, it was possible to show that cone activity never reduced the matching luminance as much as predicted by the additivity hypothesis, and sometimes even increased it. These findings suggest that cone activity can suppress brightness signals from rods.     

TRANSITION FROM ROD TO CONE VISION I. Simultaneous contrast

S tabell , B. Transition from rod to cone vision. I. Simultaneous contrast. Scand. J. Psychol ., 1969, 10 , 61–64.—Above the specific threshold, color quality depended on the test filter, while at lower intensities the same color was observed irrespectively of the test filter used, confirming the assumption that colors within the photochromatic interval are triggered by rod activity. The lawful relation between rise of specific threshold and increase of rod sensitivity was not found under the condition of simultaneous contrast.     

Spare the rod and spoil the icon

Short-term visual storage was investigated with a successive field paradigm, so that correct performance depended upon combining visual information from two targets that were never on simultaneously. In the first two experiments, the stimuli consisted of two slides, each containing a 10' red dot on a gray surround, and separated by an interstimulus interval (ISI) from 20 to 400 msec. Subjects had to determine if the dots were vertically or horizontally aligned. In Experiment 1, the stimuli had either no contrast for the rods or no luminance contrast for the cones or high contrast. At short ISIs the cone contrast determined performance, whereas at long ISIs the rod contrast determined performance. In fact, when the dots were invisible to the rods, the task was impossible for long ISIs. In Experiment 2, performance was compared for zero log rod contrast and for small departures from zero. Even a small departure from zero log rod contrast resulted in above-chance performance. In Experiment 3, the stimuli were luminous rectangles and the task was to decide whether or not a 4' spatial gap was present between the two successively presented rectangles. Wavelength, luminance, and ISI were varied under both photopic and scotopic adapting conditions. The result was that the rods performed the task for ISIs of 150 msec or longer under scotopic conditions and under the photopic conditions that we were able to test. The results of the experiments taken together are consistent with the hypothesis that the cone icon is short, whereas the rod icon is robust and long lasting.     

COLOR THRESHOLD MEASUREMENTS IN SCOTOPIC VISION: Test-stimulation varied

S tabell , U. & S tabell , B. Color threshold measurements in scotopic vision. II. Test-stimulation varied. Scand. J. Psychol ., 1968, 9, 129–132.—The color threshold curve generally coincides with the dark adaptation curve of the rods, irrespectively of test-stimulation variation, confirming the assumption that a threshold response of rods may initiate a color-related process. Variation of color threshold intensity is thus assumed to reflect variation of rod threshold intensity.     

TRANSITION FROM ROD TO CONE VISION: II. Successive contrast

S tabell , B. & S tabell , U. Transition from rod to cone vision. II. Successive contrast. Scad J. Psychol., 1969, 10 137–139—Transition from chromatic rod to chromatic cone activity during dark adaptation is interpreted as a kind of color mixing.     

The equivalent background of bleaching

Stiles and Crawford proposed that a retinal region bleached by preexposure to intense light behaves as if it were illuminated by some steady veiling or background luminance. We test this notion by comparing the afterimage of a bleaching light with a steady (and retinally stabilized) light of adjustable intensity, in the manner of Barlow and Sparrock. With their matching procedure, and also with a new procedure, we find as they did that during the rod phase of recovery the afterimage does look like a stabilized field of an intensity which, presented as a background, brings visual sensitivity to the same level. It is as if the two conditions produce equal signals at some stage of the visual pathway. Liked Barlow and Sparrock we observe a rod-cone break in the afterimage matches. However, we argue that the appearance of the rod-cone break presents a paradox and we show a way to resolve it.     

Temporal summation of pain from radiant stimulation

Latency thresholds of pricking pain, using radiant thermal stimulation, were obtained. Ten Ss were tested for 10 sessions, and, in each session, single latency determinations were made at each of 10 stimulus intensities. Each intensity was administered to a different spot along the volar surface of S’s nonpreferred forearm. Thus, 10 latency thresholds were obtained from each S at. each intensity and each spot. Initial skin temperature was controlled so that a threshold determination was made when skin temperature measured 33.5° ± 0.5° C. Analysis of variance of the log₁₀ t (seconds) values revealed a highly significant linearity of regression of log t on log intensity, thus confirming the hypothesized inverse and exponential relationship between latency and intensity. The parameters (slope and x-intercept) of the curve were discussed. The x-intercept may be interpreted as an index of an aversive threshold and could be used as a possible measure of the physiological component of pain.     

Pathways in type-B (U-shaped) metacontrast

Rod and cone targets were crossed, in every combination, with rod and cone masks in flanking-bars metacontrast. Strong type-B (U-shaped) metacontrast was obtained in each condition, contrary to the claim that rod and cone masking are independent. In each condition, visibility declined steadily with stimulus-onset asynchrony (SOA) in trials in which target and mask appeared to be simultaneous, and increased with SOA in trials in which they appeared to be successive. The 'U' results from collapsing across these different types of trials, which may reflect distinct monotonic processes in masking. Under the light adaptation conditions used the time, Tmax, at which metacontrast was at a maximum was delayed by about 25 ms if rods, rather than cones, detected the target. Whether rods or cones detected the mask hardly altered Tmax.     

Rod suppression of cone-mediated information about colour and form during dark adaptation

Following substantial bleaches, the specific form and hue thresholds were measured during dark adaptation with a test stimulus of 1 x 2 degrees at 40 degrees extrafoveally. The wavelength of the test field was varied between runs. The results show that both thresholds started to rise at about the cone-rod break of the dark-adaptation curve, irrespective of wavelength used in the test. Furthermore, the specific threshold for form was found to rise when a scotopic stimulus was superimposed on a photopic test flash. On the other hand, both thresholds remained at the cone-plateau level when the test flash was confined within the rod-free fovea. In order to explain the rise in the specific thresholds, it is suggested that signals from rods generated directly in response to the test stimulus may suppress both cone-mediated form and colour. It is also suggested that this type of rod-cone interaction represents a general characteristic involved in several kinds of visual information processing.