Effects Of Light Adaptation Research Articles

The phosphoinositide signaling cascade is involved in photoreception in the leech Hirudo medicinalis.

The effects of BAPTA, heparin, and neomycin on electrical light responses were studied in the photoreceptors of Hirudo medicinalis. Light activation produces a fast increase in intracellular Ca2+ concentration (Cai) as detected with the fluorescent Ca2+ indicator calcium green-5N. Chelating intracellular calcium by injections of 10 mmol(-1) BAPTA suppresses spontaneous quantum bumps, reduces light sensitivity by more than 2 log(10) units, and substantially increases the latent period of light responses. BAPTA strongly inhibits the plateau phase of responses to long steps of light. Injections of 45-100 mg ml(-1) of heparin act in a similar manner to BAPTA, affecting the latency of the light responses even more. De-N-sulfated heparin, an inactive analog, is almost ineffective at the same concentration compared with heparin. Heparin diminishes the light-induced Cai elevation significantly, whereas de-N-sulfated heparin does not. Intracellular injections of 50-100 mmol l(-1) of the aminoglycoside neomycin, which inhibits phospholipase-C-mediated inositol 1,4,5-trisphosphate formation, acts similar to BAPTA and heparin. Pressure injections of the hydrolysis resistant analog of inositol 1,4,5-trisphosphate, inositol 2,4,5-trisphosphate, strongly depolarize leech photoreceptors and mimic an effect of light adaptation. These results suggest a close similarity between phototransduction mechanisms in leech photoreceptors and existing models for visual transduction in other invertebrate microvillar photoreceptors.

Light-adaptive role of nitric oxide in the outer retina of lower vertebrates: a brief review.

The role of nitric oxide (NO) as a novel neurochemical mechanism controlling light adaptation of the outer retina is discussed by considering mainly published results. The emphasis is on the retinae of fishes and amphibia, but some data from the mammalian (rabbit) retinae have also been included for completeness. In the fish retina, application of NO donors in the dark caused light-adaptive photomechanical movements of cones. The normal effect of light adaptation in inducing cone contractions was suppressed by pretreatment of retinae with an NO scavenger. NO donors modulated horizontal cell activity by uncoupling the cells' lateral gap junctional interconnections and enhancing negative feedback to cones, again consistent with a light-adaptive role of NO. Direct evidence for light adaptation-induced release of NO has been obtained in fish (carp) and rabbit retinae. The results strongly suggest that control of retinal light adaptation is, under multiple neurochemical control, with NO and dopamine having an interactive role.

Transient enhancing of cone electroretinograms following exposure to brighter photopic backgrounds

Normal subjects were first light adapted to a standard photopic background and a control photopic ERG was obtained. The subjects were then light adapted to a brighter background for 5 min at the end of which the luminance was returned to the control background and ERGs were taken at regular intervals. Most of the ERG/OP components were transiently enhanced following the above procedure. Given that the previously reported photopic light adaptation effect occurred following an increase in the luminance of the adapting field (from dark adaptation to light adaptation) while that reported in the present study is triggered following a decrease in the level of light adaptation, the opposite effects noted might suggest that the two retinal events result from the same, not yet identified, cone adaptation mechanisms which are solicited in an opposite way.

GABACReceptors Control Adaptive Changes in a Glycinergic Inhibitory Pathway in Salamander Retina

We studied the role of GABA in adaptive changes in a lateral inhibitory system in the tiger salamander retina. In dark-adapted retinal slice preparations picrotoxin caused a slow enhancement of glycine-mediated IPSCs in ganglion cells. The enhancement of glycinergic IPSCs developed slowly over the course of 5-20 min, even though picrotoxin blocked both GABA(A) and GABA(C) receptors within a few seconds. The slow enhancement of glycinergic IPSCs by picrotoxin was much weaker in light-adapted preparations. The slow enhancement of glycinergic inhibitory inputs was not produced by bicuculline, indicating that it involved GABA(C) receptors. The responses of ganglion cells to direct application of glycine were not enhanced by picrotoxin, indicating that the enhancement was not caused by an action on glycine receptors. In dark-adapted eyecup preparations picrotoxin caused a slow enhancement of glycinergic IPSPs and transient lateral inhibition produced by a rotating windmill pattern, similar to the effect of light adaptation. The results suggest that the glycinergic inhibitory inputs are modulated by an unknown substance whose synthesis and/or release is inhibited in dark-adapted retinas by GABA acting at GABA(C) receptors.

Effects of nitric oxide, light adaptation and APB on spectral characteristics of H1 horizontal cells in carp retina.

The spectral characteristics of cone-driven horizontal cells of H1 subtype (H1 HCs) receiving main synaptic input from red-sensitive cones were studied in light- and dark-adapted retinae of carp. The spectral sensitivity profile of H1 HCs in dark-adapted retinae was practically the same as the absorption spectrum of red-sensitive cones. Light-adaptation decreased the sensitivity preferentially in the short-wavelength (blue/green) region, resulting in a relative enhancement of the 617 nm peak. Application of nitric oxide (NO) donors, sodium nitroprusside (SNP) and S-nitrosoglutathione (SNOG or GSNO), or dopamine to dark-adapted retinae decreased the sensitivity preferentially in blue/green region, an effect similar to that of light-adaptation. Application of haemoglobin (Hb, an NO scavenger) or 2-amino-4-phosphonobutyrate (APB, a metabotropic glutamate receptor agonist), to light-adapted retinae increased the sensitivity preferentially in the blue/green region, an effect similar to dark-adaptation. The photoresponses of H1 HCs were univariant in dark-adapted retinae as well as Hb-treated light-adapted retinae. In light-adapted retinae with normal Ringer, however, the univariance did not hold. These results suggested that the photoresponses of H1 HCs to short-wavelength stimuli contain a depolarising (sign-reversing) component, which can be activated by light-adaptation or application of NO and dopamine, and inactivated by dark-adaptation or deprivation of NO or application of APB.

Adaptation-dependent plasticity of rod bipolar cell axon terminal morphology in the rat retina.

We chose synaptic terminals of rat rod bipolar cells as a model system to study activity-related changes in the overall morphology and the fine structure of synaptic sites. Using confocal laser scanning microscopy in conjunction with three-dimensional reconstruction and electron microscopy, we examined the effect of light and dark adaptation on axon terminals identified by protein kinase C (PKC) immunoreactivity. Rod bipolar cell axon terminals consisted of 2-3 polymorphic boutons situated close to the ganglion cell layer and a single ovoid swelling located more distally. Both components of the terminal complex showed adaptation-dependent differences in the distribution of PKC immunoreactivity and in their morphology. In light-adapted rod bipolar cell axon terminals, PKC immunoreactivity was homogeneously distributed throughout the cytoplasm, whereas terminals from dark-adapted animals showed PKC immunoreactivity preferentially localised in the submembrane compartment and a reduced staining of the more central cytoplasm. In three-dimensional reconstructions of optical sections and at the ultrastructural level, the shape of light-adapted axon terminals was round and smooth and exhibited more convexly curved synaptic membranes. In contrast, dark-adapted terminals had irregular contours, numerous dimples and a concave synaptic curvature. No spinules of bipolar cell terminals were observed in dark-adapted material. These observations are discussed in the context of activity-related morphological plasticity of central nervous system synapses and of the functions of PKC in the cycle of vesicle fusion and retrieval at the tonically active ribbon synapses of the rod bipolar axon terminal.

Adaptation-dependent changes of bipolar cell terminals in fish retina: Effects on overall morphology and spinule formation in Ma and Mb cells

We have investigated the effects of light and dark adaptation on the overall morphology of bipolar cell (BC) terminals in sublaminae a and b of the inner plexiform layer after labelling with Lucifer Yellow (LY) and PKC immunostaining using confocal laser scanning microscopy and serially sectioned material for electron microscopy. Three-dimensional reconstructed terminals showed marked adaptation-dependent changes of their morphology. Terminals of mixed rod-cone BCs in sublamina a (Ma BC) were irregular and scalloped in light adapted, but smooth and regular in dark-adapted specimens. Terminals from mixed rod-cone BCs in sublamina b (Mb BCs) exhibited an opposite behaviour. At the ultrastructural level, bipolar terminals in both sublaminae showed fingerlike extensions (spinules) invaginating presynaptic amacrine cell (AC) processes. Sixty-two percent of the dark-adapted Mb terminals in sublamina b showed spinules, whereas 21% of the light-adapted terminals had spinules. By contrast, 50.6% of the light-adapted Ma terminals in sublamina a formed spinules, compared to 17.8% of the dark-adapted Ma terminals in this sublamina. These observations reflect the functional subdivision of the inner plexiform layer in an inner ON- and an outer OFF-centre lamina. Our findings suggest that the synaptic plasticity of BC axon terminals may be due to differences of BC membrane potential, or the activity of AC input onto bipolar terminals. They may contribute to processes of fine tuning regulating the efficiency of AC-3-BC interaction under varying adaptation conditions.

Spectral plasticity of H1 horizontal cells in carp retina: independent modulation by dopamine and light-adaptation.

It was shown previously that the spectral sensitivity of luminosity/H1-type horizontal cells (HCs) in carp retinae reflects the absorption spectrum of red-sensitive cones for long wavelengths but can appear highly variable and "truncated' in the short-wavelength region of the spectrum. We have found that light-adaptation sharpened the red-sensitive spectral peak and decreased the blue/red response amplitude ratio (B/R ratio), mainly by decreasing the response to short-wavelength stimuli. The adaptation effect was more pronounced for red background light than for blue. During dark adaptation, the B/R ratio increased steadily. Exogenous dopamine (DA; 5 microM) changed the spectral response profile in a similar way to light-adaptation. However, the effect of light-adaptation in reducing the B/R ratio was still seen in retinae bathed in 5 microM DA. This effect of background adaptation was also recorded in retinae bathed in 37 microM haloperidol, as well as in retinae pretreated with 6-hydroxydopamine (i.e. DA-depleted). The results suggest that (i) short-wavelength-sensitive cones play a dynamic role in determining the spectral response profile of H1 HCs and (ii) spectral response characteristics are modulated independently by exogenous DA and an unknown endogenous neuromodulator which is activated by light-adaptation.

Contrast discrimination at high contrasts reveals the influence of local light adaptation on contrast processing

Previous measurements of contrast discrimination threshold, Δ C, as a function of pedestal contrast, C, for sine-wave gratings have shown a power law relationship between Δ C and C at suprathreshold levels of C. However, these studies have rarely used contrasts greater than 50%. Whittle (1986), using incremental and decremental patches, found that Δ C increased with C only up to about 50%. At higher contrasts it decreased. Since a periodic stimulus can be considered to be composed of increments and decrements, we thought we might find such an inverse U-shaped function for gratings if we used contrasts up to 100%. We tested this for both sine-wave and square-wave stimuli at spatial frequencies from 0.0625 to 8.0 c/deg. We found that for frequencies up to 0.5 c/deg, Δ C in nearly all cases ‘dipped down’ after about C = 50% contrast. At 4.0 and 8.0 c/deg, however, no dip-down occurred. Additional experiments showed that the dip-down was unlikely to be due to cortical long-term adaptation and most likely an effect of localized light adaptation to the dark bars. We argue that the absence of dip-down at high spatial frequencies was mainly due to the attenuation of contrast by the optics of the eye. As for the results of Whittle (1986), a Weber's Law in W = ( L max − L min) L min describes the inverse U-shaped contrast discrimination function well. Two other contrast expressions also linearize the data on log-log plots. We show how some familiar notions about the physiological operation of localized light adaptation can easily account for the form of the contrast discrimination function. Finally we estimate the number of discriminable steps in contrast from detection threshold to maximum contrast for the various spatial frequencies tested.

Localization of heme oxygenase in rat retina: effect of light adaptation

Heme oxygenase-2 isozyme is the predominant form of heme oxygenase in rat brain by western blot analysis. Heme oxygenase-1 isozyme is not induced by light adaptation in rat retina by western blot analysis. Immunocytochemistry localizes heme oxygenase-2 in three areas of the retina: the retinal pigment epithelium, inner segment and external nuclear layers of the rat retina. Ganglion cells and cell bodies of the internal nuclear layer of the retina and Müller cells were largely unstained for heme oxygenase-2. The localization of heme oxygenase-2 in the retina implies that its function is not associated with phototransduction. Also, light adaptation does not appear to induce heme oxygenase; 1, a measure of oxidative injury.

Light adaptation of the human photopic oscillatory potentials: influence of the length of the dark adaptation period.

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.

Light adaptation and the rising phase of the flash photocurrent of salamander retinal rods.

1. Both theory and analysis of photocurrents in retinal rods show that phosphodiesterase activity after a flash rises initially as a delayed ramp. 2. The effect of light adaptation on the flash-induced rise in phosphodiesterase activity deduced from photocurrent responses was investigated. 3. Background adaptation reduces the deduced rate of rise of phosphodiesterase activity. The effect is most prominent for bright backgrounds and moderate flashes. There is little reduction for bright flashes, even in bright backgrounds. There is no effect for weak backgrounds. 4. Light adaptation after bleaching visual pigment produces a reduction in the deduced rise of phosphodiesterase activity for all flashes. For bright flashes, the reduction is explained by the reduction in quantum catch. For moderate flashes, there is an extra reduction, similar to the reduction produced by the equivalent background. 5. The results provide support for the idea that a reduction in the amplification step of phototransduction functions as part of the mechanism of light adaptation in rods. The dependence on flash intensity of the background-induced reduction in phosphodiesterase activation could imply a feedback mechanism on the activation steps of phototransduction.

Modulation of transduction gain in light adaptation of retinal rods.

The effect of light adaptation on the period of photocurrent saturation induced by a bright stimulating flash was examined in rod photoreceptors of the larval-stage tiger salamander (Ambystoma tigrinum). Using suction electrodes, photocurrent responses to brief flashes were recorded from single, isolated rods in the presence and absence of steady background illumination. Background light decreased the saturation period (T) measured at fixed flash intensity (fixed If) and in this respect light-adapted the saturating response. Effects of the background on responses to weak (i.e. subsaturating) and bright flashes were compared with changes in a parameter, phi = e-delta T/TR*, where delta T is the decrease in saturation period, and where TR* is the slope of the line that relates T and ln If in a given state of adaptation. Dark- and light-adapted responses to flash intensities IDf and ILf, respectively, exhibited similar absolute peak photocurrent and falling-phase kinetics when IDf and ILf satisfied the relation, IDf = phi (ILf + IbTR*), where Ib is the background intensity. It is argued that phi approximates the relative PDE*/R* gain of transduction, i.e. the relative peak level of activated cGMP phosphodiesterase (PDE*) produced by a given, small amount of photoactivated visual pigment (R*). Interpreted on this view, the results imply that light adaptation derives largely from a decrease in PDE*/R gain, rather than from the stimulation of guanylate cyclase activity. The data are consistent with the possibility that modulation of the lifetime of PDE* underlies the background dependence of phi.

Controlling Unwanted Sources of Threshold Change in Disability Glare Studies

One of the main purposes of cataract surgery is to reduce the debilitating effects of light scatter by the lens. An important effect of this intraocular forward scatter is to produce a veiling retinal illuminance from a glare source. The retinal contrast of the stimulus is thus reduced, and a quantifiable "disability glare" effect may be measured. A complication is introduced by the fact that intraocular scatter arises from other sources in addition to the lens. All present tests of glare disability fail to eliminate or control for these factors, nor do they control for the light adaptation effect of the glare source. The effects of lens removal may not be predicted accurately by such tests, and patients may report little improvement in visual function after surgery. The goal of this research is to provide the background for developing a method of measuring disability glare due only to light scatter in the lens. Thirty-seven patients with mild to moderate cortical cataracts served as subjects. A disability glare effect (DG) was measured with a Maxwellian-view system that eliminates or controls scatter from the iris, sclera, and retina, and which controls for light-adaptation effects from the glare source. DG with this method showed a significant correlation with objective measures of lens light back scatter. Suggestions were made for improvements in the apparatus and procedures for future clinical studies.

Decreased energy requirement of toad retina during light adaptation as demonstrated by 31P nuclear magnetic resonance.

1. The effect of light and dark adaptation on the levels of phosphorus metabolites (nucleotide di- and triphosphates, phosphocreatine, pyridine nucleotide, inorganic phosphate, phosphodiesters, phosphomonoesters, and uridine diphosphate-glucose) in the toad (Bufo marinus) retina and retinal extracts was studied by 31P nuclear magnetic resonance (NMR) spectroscopy. 2. Spectra were acquired using an NMR probe specifically designed for superfusion and illumination of a single retina. Retinae were maintained at a steady state for up to 10 h in an electrolyte solution containing 10 mM Hepes buffer and bubbled with 98% O2-2% CO2, pH 7.8 at 20 degrees C. 3. The intracellular concentrations of the phosphorus metabolites were measured in total darkness or during prolonged exposure to light. The concentration of nucleoside triphosphates (NTP) in the dark-adapted retina was about 1.5 mM and that of phosphocreatine (PCr) was about 0.7 mM. 4. In saturating levels of light, 6.0 x 10(11) or 1.5 x 10(13) quanta s-1 cm-2 at 520 nm, the levels of PCr and phosphomonoesters rose, the levels of NTP and protons (pH) were maintained, and the levels of pyridine nucleotides and nucleotide diphosphates (NDP) fell. 5. A rise in the level of PCr in the presence of an unchanged level of NTP in the light-adapted retina indicates that the energy consumption of the retina is greater in the dark. 6. These results are in agreement with the results of oxygen consumption, glucose dependence, and electrophysiological studies which also indicate that the metabolic energy requirement of the retina decreases in light.

Light-dependent plasticity of the synaptic terminals of Mb bipolar cells in goldfish retina.

We recently described spine-like protrusions (spinules) from the synaptic terminals of mixed rod-cone (Mb) bipolar cells of goldfish retina that invaginated about 5% of the presynaptic amacrine cell processes (Yazulla and Studholme, J Comp Neurol 310:11, 1991). In view of reports of a light/dark dependent-plasticity on the formation of dendritic spinules on goldfish horizontal cells (Raynauld et al., Science 204:1436, 1979; Wagner, J Neurocytol 9:573, 1980), we investigated the possibility that Mb terminal spinules also might show light/dark-dependent plasticity. Retinas were obtained at noon time from light-adapted and 3-hour dark-adapted goldfish and processed for electron microscopy using conventional histological procedures and by preembedding immunocytochemistry to detect protein kinase C immunoreactivity. Two effects of light adaptation on Mb terminal morphology were observed. First, the surface of Mb terminals was significantly more irregular after dark adaptation than light adaptation. With dark adaptation, Mb terminals appeared "amoeboid," with numerous cytoplasmic extensions between the presynaptic processes. Second, spinules were sevenfold more frequent after dark adaptation than light adaptation (8% vs. 1.2% of the presynaptic processes were invaginated by spinules). We suggest that the increased frequency of spinules during dark adaptation is related to an enhancement of synaptic transmission from a minor amacrine cell input when the major input from GABAergic amacrine cells is reduced. Also, the irregular surface of dark-adapted Mb terminals may be related to the reduction of synaptic input during dark adaptation.

The Effect of Light Adaptation on Scotopic Spatial Summation in 10-Week-old Infants

Psychophysical area-intensity functions of individual 10-week-old human infants and adults were obtained in the dark adapted state, and in the presence of a steady background that elevated threshold 1 log unit above the dark adapted level. For dark adapted infants, the mean diameter for complete spatial summation (4.42°; SD: 1.67°) was significantly larger than that of adults (2.32°; SD: 0.09°). The background reduced the mean critical diameter to 2.67° for infants (SD: 0.64°) and to 1.16° for adults (SD: 0.08°). Spatial probability summation has similar effects on infant and adult thresholds, and, therefore, does not appear to account for the developmental decrease in critical diameters. Rather, decreases in receptive field size are suspected.

Electroretinographic Monitoring of Retinal Function During Eye Surgery

We performed a pilot study of electroretinographic monitoring of retinal function during eye surgery performed with local anesthesia. A contact lens with a built-in light-emitting diode was sterilized and used as both a stimulus source and a recording electrode for 30-Hz flicker electroretinograms. Each recording required 7 seconds, during which the operating light (but not the room light) was switched off. Electroretinograms recorded during surgery on 10 patients with minimal retinal abnormality were evaluated for the effects of light adaptation caused by regular exposure to an operating light source or operating microscope. The fluctuations of the amplitude and peak time were small, indicating that this technique can be used to monitor retinal function during eye surgery. Some results during retinal and vitreous surgeries are shown.

Effects of light adaptation on the response characteristics of human oscillatory potentials

We have examined the response characteristics of the oscillatory potentials (OPs) of the human electroretinogram (ERG) obtained to ganzfeld flash stimuli presented against adapting fields. First, we determined the extent to which the OPs obtained to high luminance flashes change during the course of light adaptation to a cone-isolating adapting field. Regardless of the number of OP wavelets, the last OP wavelet increased in amplitude and decreased in implicit time to a greater extent than did the earlier wavelet(s). In addition, we examined the role of both flash and adapting field luminance in determining the wave form of the OPs. For each adapting field luminance that was tested, the number of OP wavelets increased as flash luminance increased, primarily resulting from the splitting of the last OP into 2 distinct wavelets. While the number of OP wavelets generally decreased as adapting field luminance increased, the amplitude of the last OP became larger. These functional distinctions between the last and the earlier wavelets are consistent with their representing the activity of different retinal generators.

Gap junction particle density of horizontal cells in goldfish retinas lesioned with 6‐OHDA

The I1 dopaminergic interplexiform cells of the fish retina are believed to modulate horizontal cell coupling by increasing gap junction resistance. Dopamine also modulates the morphology of horizontal cell gap junctions and mimics the effects of light adaptation. To determine whether the light-dependent changes in gap junction morphology are due to endogenous dopamine release, horizontal cell gap junctions were studied in goldfish retinas lacking dopaminergic neurons. Dopaminergic interplexiform cells were destroyed by intraocular injections of 6-hydroxydopamine in both eyes. After lesioning, fish were treated in one of four ways: (1) light-adapted, (2) dark-adapted (1 hour), (3) light-adapted and given an intraocular injection of dopamine, or (4) dark-adapted (1 hour) and injected with dopamine. The effectiveness of lesioning was evaluated by autoradiographic detection of [3H]-dopamine uptake in the retina of one eye. Retinas in which lesioning of the contralateral eye was deemed effective were processed for freeze-fracture electron microscopy and the particle density of horizontal cell gap junctions determined. Lesioned retinas, whether light- or dark-adapted, had elevated horizontal cell soma gap junction particle densities compared to lesioned retinas treated with dopamine. These results demonstrate that high soma gap junction particle densities can be correlated with the absence of dopamine and low densities associated with the presence of dopamine. The differences in gap junction particle density between lesioned and lesioned + dopamine-treatment were similar to differences between nonlesioned dark-adapted (1 hour) and light-adapted retinas, respectively. Therefore, the particle density of light- and dark-adapted soma gap junctions suggests a greater release of dopamine in light-adapted fish than in 1 hour dark-adapted fish.(ABSTRACT TRUNCATED AT 250 WORDS)