[Dark adaptation in patients with chronic liver diseases. 1. Disturbances in dark adaptation in patients with chronic liver].
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Dark-adapted adults' electroretinographic b-wave thresholds are approximately 2 log units below a-wave thresholds and approximately 3 log units above the perceptual threshold, and their perceptual and the visually evoked cortical potential (VEP) thresholds are similar. Dark-adapted infants' scotopic a- and b-wave thresholds for full-field stimuli are both about 0.5 log units above those of adults, but their scotopic VEP thresholds for such stimuli have not been studied. We obtained scotopic VEP thresholds for brief, full-field stimuli from dark-adapted, infants and adults to consider the relationships of the cortical responses to the responses of more distal structures, namely the rod photoreceptors (scotopic a-wave) and ON-bipolars (scotopic b-wave). The median VEP threshold of infants is 0.5 log unit above that of adults. Thus, the relationships of a- and b-wave and VEP thresholds in infants are similar to those in adults. These results are consistent with rod cell immaturities being the primary determinant of the difference between infants' and adults' thresholds.
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Nanosecond time-resolved absorption spectra have been measured throughout the photocycle of bacteriorhodopsin in both light-adapted and dark-adapted purple membrane (PM). The data from dark-adapted samples are interpretable as the superposition of two photocycles arising independently from the all-trans and 13-cis retinal isomers that coexist in the dark-adapted state. The presence of a photocycle in dark-adapted PM which is indistinguishable from that observed for light-adapted PM under the same experimental conditions is demonstrated by the observation of the same five relaxation rates associated with essentially identical changes in the photoproduct spectra. This cycle is attributed to the all-trans component. The cycle of the 13-cis component is revealed by scaling the data measured for the light-adapted sample and subtracting it from the data on the dark-adapted mixture. At times less than 1 ms, the resulting difference spectra are nearly time-independent. The peak of the difference spectrum is near 600 nm, although there appears to be a slight (approximately 2 nm) blue-shift in the first few microseconds. Subsequently the amplitude of this spectrum decays and the peak of the difference spectrum shifts in two relaxations. Most of the amplitude of the photoproduct difference spectrum (approximately 80%) decays in a single relaxation having a time constant of approximately 35 ms. The difference spectrum remaining after this relaxation peaks at approximately 590 nm and is indistinguishable from the classical light-dark difference spectrum, which we find, in experiments performed on a much longer time scale, to peak at 588 nm. The decay of this remaining photo-product is not resolvable in the nanosecond kinetic experiments, but dark adaptation of a completely light-adapted sample is found to occur exponentially with a relaxation time of approximately 2,000 s under the conditions of our experiments.
In teleosts, the basal part of the retinal pigment epithelium (RPE) is relatively smooth, i. e., it is free of basal membrane infoldings. In the featherback, Notopterus notopterus, whereas this is the situation in light adaptation, during dark-adaptation, especially when kept for prolonged periods (6-9 hour), numerous infoldings appear at the basal region, as found uniquely by transmission electron microscopy. In this teleost, during retinomotor movements, the rods move vitreally during dark-adaptation, while the cones do not elongate, and remain stationary in both light- and dark-adaptation. The significance of the appearance of basal infoldings in dark-adapted RPE is explained in terms of the pattern of retinomotor responses and the features of RPE and photoreceptors in this species. It is suggested that (1) the thick, impervious tapetal layer present in the RPE, (2) the unusual position of the photoreceptors in the visual cell layer of dark-adapted retina, and (3) the presumably high demand for glucose and O2 of the outer retina during dark-adaptation might contribute to cause this phenomenon in this species. The available evidence tend to associate this phenomenon with the involvement of the RPE in nutrient and O2 delivery to the photoreceptors via the basal infoldings of the RPE in dark-adapted state in this species. This has not been reported for any other teleosts to date.
The effects of dark adaptation on the response properties of ganglion cells have been documented extensively in the cat retina. To pinpoint the different retinal mechanisms that underlie these effects, we studied the response characteristics of cat horizontal (H) cells during prolonged dark adaptation. H-cell responses were recorded intracellularly in the optically intact, in vivo eye. To disentangle rod and cone contributions, sensitivity changes during dark adaptation were tracked with white light and with monochromatic lights that favored either rod or cone excitation. Stable, long-lasting recordings allowed us to measure changes of sensitivity for adaptation periods up to 45 min. Thresholds for white light and 503-nm monochromatic light decreased steadily and in parallel. The maximum increase of sensitivity, after extinguishing a photopic adaptation light, was 1.8 log units only, reached after about 35 min. Sensitivity for 581-nm lights also increased steadily, but at a shallower slope. The steady increase of sensitivity was concomitant with a linear shift in resting membrane potential and with an increase in relative rod contribution to the threshold responses. Even though small-amplitude responses were rod dominated after prolonged dark adaptation, sensitivity to rod signals remained relatively low, compared to sensitivity of cone responses or to the absolute sensitivity of ganglion cells. This suggests that the cone-H-cell pathway plays no role in the dark-adapted cat retina.
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The recovery of rod responsiveness after saturating flashes is greatly retarded above a certain critical level of rhodopsin bleaching (approximately 0.1%). A mathematical description of the process of turn-off of the phototransduction cascade allows attributing different phases of the recovery to specific products of rhodopsin photolysis. The fast phase is determined by quenching of metarhodopsin II and activated transducin. The slow phase is controlled by decay of partially inactivated (phosphorylated and arrestin-bound) metarhodopsins, and by regeneration of rhodopsin. The transition between the two regimes of adaptation is rather abrupt, occurring within a few-fold range of stimulus intensity. This marks the border between reversal of light adaptation and dark adaptation, as it is commonly defined.
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Composition of retinal isomers in three proton pumps (bacteriorhodopsin, archaerhodopsin-1, and archaerhodopsin-2) was determined by high performance liquid chromatography in their light-adapted and dark-adapted states. In the light-adapted state, more than 95% of the retinal in all three proton pumps were in the all-trans configuration. In the dark-adapted state, there were only two retinal isomers, all-trans and 13-cis, in the ratio of all-trans: 13-cis = 1:2 for bacteriorhodopsin, 1:1 for archaerhodopsin-1, and 3:1 for archaerhodopsin-2. The difference in the final isomer ratios in the dark-adapted bacteriorhodopsin and archaerhodopsin-2 was ascribed to the methionine-145 in bacteriorhodopsin. This is the only amino acid in the retinal pocket that is substituted by phenylalanine in archaerhodopsin-2. The bacteriorhodopsin point-mutated at this position to phenylalanine dramatically altered the final isomer ratio from 1:2 to 3:1 in the dark-adapted state. This point mutation also caused a 10 nm blue-shift of the adsorption spectrum, which is similar to the shift of archaerhodopsin-2 relative to the spectra of bacteriorhodopsin and archaerhodopsin-1.
Dark-adapted electroretinograms were obtained over a 3.6-log range of stimulus intensities from 17 black and 15 white normal subjects. Subjects were grouped on the basis of light or dark fundus pigmentation, determined from digitized fundus photographs. B-wave amplitudes for each group were fitted by the Naka-Rushton equation, and the measures Vmax, log K, and n were determined. The luminance-response functions revealed that subjects with light fundi had larger b-wave amplitudes at all luminance levels. There was a significant difference between groups for Vmax and n but not for log K. A comparison of b-wave implicit times showed no significant difference between subjects with dark and light fundi. Ancillary tests and multiple regression analysis suggested that the relationship between Vmax and fundus pigmentation could not be attributed to age, gender, refractive error, axial length or intraocular pressure. The results have implications for the collection of normative electroretinographic data and for the interpretation of electroretinogram results.
Light and dark adaptation of the phototropism of Phycomyces sporangiophores were analyzed in the intensity range of 10(-7)-6 W X m-2. The experiments were designed to test the validity of the Delbrück-Reichardt model of adaptation (Delbrück, M., and W. Reichardt, 1956, Cellular Mechanisms in Differentiation and Growth, 3-44), and the kinetics were measured by the phototropic delay method. We found that their model describes adequately only changes of the adaptation level after small, relatively short intensity changes. For dark adaptation, we found a biphasic decay with two time constants of b1 = 1-2 min and b2 = 6.5-10 min. The model fails for light adaptation, in which the level of adaptation can overshoot the actual intensity level before it relaxes to the new intensity. The light adaptation kinetics depend critically on the height of the applied pulse as well as the intensity range. Both these features are incompatible with the Delbrück-Reichardt model and indicate that light and dark adaptation are regulated by different mechanisms. The comparison of the dark adaptation kinetics with the time course of the dark growth response shows that Phycomyces has two adaptation mechanisms: an input adaptation, which operates for the range adjustment, and an output adaptation, which directly modulates the growth response. The analysis of four different types of behavioral mutants permitted a partial genetic dissection of the adaptation mechanism. The hypertropic strain L82 and mutants with defects in the madA gene have qualitatively the same adaptation behavior as the wild type; however, the adaptation constants are altered in these strains. Mutation of the madB gene leads to loss of the fast component of the dark adaptation kinetics and to overshooting of the light adaptation under conditions where the wild type does not overshoot. Another mutant with a defect in the madC gene shows abnormal behavior after steps up in light intensity. Since the madB and madC mutants have been associated with the receptor pigment, we infer that at least part of the adaptation process is mediated by the receptor pigment.
The average dark adaptation time for newly recruited Black mineworkers is longer than that reported for Whites. While this longer dark adaptation time may result from a number of pathological conditions, its most likely cause is a deficiency of vitamin A in the diet. A study was conducted in which the vitamin A level of the blood and the dark adaptation times were correlated in a group of Black mineworkers upon arrival and again after 4 - 6 months continuous work underground in a gold mine. The study indicated that a very marked decrease occurred in the miners' serum vitamin A level between the time of arrival at the mine and the second examination. This difference was found to be highly significant. This decrease was accompanied by an increase in mean dark adaptation time, which was significant at the 5% level. These observations accord with the findings of a previous study and are indicative of an inadequate dietary intake of vitamin A during the period when the miners were on the mine diet.
Dark incubation of envelope vesicles derived from a strain of Halobacterium halobium that lacks bacteriorhodopsin but contains halorhodopsin and a third rhodopsin-like pigment caused a decrease in the flash yield [the amplitude of a transient absorbance change of flash reactive component(s) by flash] of halorhodopsin but not the rhodopsin-like pigment. The flash yield decreased to reach a low steady level after incubation for about 4 days in the dark. The flash yield of halorhodopsin at any stage of dark incubation was increased by actinic illumination of the vesicles. The flash yield at 490 nm (absorbance increase) was found to be approximately proportional to that at 590 nm (absorbance decrease). These results indicate that halorhodopsin in the envelope vesicles has two forms, dark and light adapted, and that the halorhodopsin phototransient absorbing at 490 nm is originated from the light-adapted form. A difference spectrum between these two forms of halorhodopsin shows that the light-adapted halorhodopsin was red-shifted from the dark-adapted form. The light-induced membrane potential was measured by tetraphenylphosphonium uptake. The uptake by the dark-adapted vesicles was slower than that by the light-adapted vesicles, suggesting that only the light-adapted halorhodopsin has ion-transporting activity.
The dark-adapted cone electroretinogram (ERG) is difficult to isolate because of unwanted rod intrusion. We compare dark-adapted cone estimates derived using three techniques. The first uses the cone response on a moderate rod saturating background to estimate the dark-adapted cone response. The second uses red and blue flashes to tease apart cone and rod responses (red-minus-blue technique, [Investigative Ophthalmology and Visual Science 31 (1990) 2283]). The third uses a bright flash to temporarily saturate rods, followed by a test flash that generates a putative cone-only response (2-flash technique [Investigative Ophthalmology and Visual Science 36 (1995) 1603]). By subtracting the cone estimates from 'mixed' ERG responses in the dark, rod isolated responses can be derived. The rod phototransduction parameters, derived using a computational model, are similar using the light-adapted and 2-flash cone estimates, but differ using the red-minus-blue estimates. The 2-flash cone estimate gives a cone waveform similar to the dark-adapted response of a patient with Oguchi stationary night blindness (a patient with no rod ERG responses and normal cone ERG responses). The growth of the cone response during light adaptation to steady backgrounds causes significant differences between the light-adapted and 2-flash cone waveforms at times beyond the first few milliseconds.
Cone dark adaptation curves were measured in a rhesus monkey using the electroretinogram (ERG) response to a 40 Hz flickering stimulus. The influence of anesthetics on the time course of dark adaptation was studied. All volatile anesthetics tested (methoxyflurane, halothane, enflurane, ether, chloroform) retarded dark adaptation but to different degrees; urethane, ethyl alcohol, and Freon 11 also retarded dark adaptation. No effect was found for barbiturates and ketamine. It seems unlikely that metabolites play a role in the observed phenomena. A literature survey reveals that several studies on dark adaptation or visual pigment regeneration might have suffered from influences of the anesthetic used. The cause of the phenomenon might lie either in anesthetics-induced membrane changes or in hindrance of the isomeration of 11-trans retinal to 11-cis retinal.
In zebrafish, during dark adaptation following bright light adaptation, the dominance of electroretinogram (ERG) b- and d-waves switches. In the early dark adaptation, when visual sensitivity is cone-dominant, both the b- and d-waves are readily recorded. In the late dark adaptation, along with the increase of rod sensitivity, the b-wave becomes dominant whereas the d-wave is gradually lost. The time for the ERG b- and d-wave dominance transition varies between the day and night. The transition requires a longer amount of time in the night and early morning than in the afternoon. This pattern of timing for ERG b- and d-wave dominance transition persists in constant light and can be reversed after exposure to a reversed light-dark cycle. The data suggest that the transition of the dominance of ERG b- and d-waves is regulated by an endogenous circadian clock.
Photopic electroretinograms recorded immediately after a period of dark adaptation show a regular increase in amplitude with time spent in light. The retinal mechanisms at the origin of this light adaptation effect remain obscure. The purpose of this study was to investigate the duration of the dark adaptation period needed to produce an optimal light adaptation effect as demonstrated by photopic oscillatory potential recordings. Our results indicate that the light adaptation effect can be separated into two distinct processes. The first one, activated early in the dark adaptation process, reduces the amplitude of the fourth oscillatory potential to 32% of control after less than 5 min of dark adaptation, while the second process, activated after more than 10 min of dark adaptation, appears to impact solely the amplitude of the earlier oscillatory potentials 2 and 3. Our results suggest that the light adaptation effect is mediated by two distinct retinal pathways or mechanisms.