Differentiation of the bimodal stimuli in a frog's retina

In work electric activity of frog's retina was investigated by silent substitution technique. Electroretinogram was recorded as a response to abrupt exchange of the referent stimulus-line with fixed values of luminance and orientation to test lines with varied luminance and orientations. As a result of the analysis it has been allocated two types of responses of a retina. The response to onset-offset of a stimulus-line was similar to the response at homogeneous illumination of a retina (ERG), and was characterized by both the high amplitude of b-wave (hundreds mkV) and significant asymmetry of b- and d-waves. Whereas the same waves in response to substitution of the same stimuli were more symmetric and had on ten times smaller amplitudes. Such activity of frog's retina was referred as pattern electroretinogram (PERG) recorded in a high vertebrate's retina as response to stimuli whose contrast was temporally modulated. The analysis of interaction of luminance and line orientation channels in retina was carried out on the base of construction V-shaped functions of stimuli differentiation. It has shown, that activities of both channels are linearly summarized in PERG. It means independent and parallel functioning of these mechanisms. However, it takes the short subdivision of luminance, namely, when luminance of test line not far from luminance of referent line. At the same time, from the moment of the double prevalence of test line in relation to referent line, growth of PERG amplitude has nonlinearly form. Such two-stage changing of PERG amplitude speaks presence in a retina of a frog of two mechanisms of coding of luminance. One mechanism coding light intensity by power of the discharge, it forms the information on an absolute level of light in the environment. Its activity is caused basically, by receptors and cells of external plexiform layer and is submitted by b-wave of electroretinogram. Other mechanism submitted in PERG, is based on the vector code of stimulus, it forms the information on spatial and time differentiation of a light in the visual field and is connected, basically, with cells of internal plexiform layer of frog's retina.     

Significance of presetting an analyzer system in discriminating light intensity in the cat

EEG and behavioural reactions were studied at the action of signal and nonsignal light flashes. The minimal time of stimulus exposition necessary for preserving differences in EEG activation reactions to nonreinforced stimuli of different intensity (in a diapason from 3.5 to 250 lk) was shown to exceed 1 s. After giving the signal meaning to the flashes the minimal time of stimulus exposition when the animals were capable to discriminate intensity of the flashes (in a diapason from 3.5 to 30 lk) just at the first presentation, was equal to 0.1 s. Decrease of the minimal time of stimulus exposition necessary for discrimination of the light signals was supposed to be stipulated by their biological significance and presetting of the analyzing system being organized by preliminary learning a crude analysis of stimuli. Discrimination of the light stimuli of short duration was impossible without such presetting.     

Chromatic component of the frog electroretinogram

To single out the chromatic constituent of frog electroretinogram the method of great (superthreshold) differences between stimuli is used. The stimuli are presented as instantly replaced colour flashes. It is shown that the amplitude of the b-wave of the electroretinogram recorded during the replacement of the equally bright stimuli is determined only by their colour difference.     

The behavior of the b-wave in the electroretinogram of light-adapted frogs during serial light exposure]

In light-adapted frogs (adaptation luminance 10(3) asb) repeated flashes of high intensity (10(5) asb) caused decrease of the b-wave in the electroretinogram as compared to the results obtained with dark-adapted animals. The amplitude changes were monotonous in all cases; they were dependent on the spacing and duration of the stimulus. With short intervals (2-10 sec) the amplitude adjusted to a new level no later than after 8 flashes. With longer intervals (30 and 60 sec), however, the amplitude decreased slowly and steadily over the whole period of experiment. Under these conditions, adjustment to a new level was prevented by disturbing factors, which additionally reduced the retinal sensitivity depending on the duration of the experiment.     

Comparison of Stimulus Energies Required to Elicit the ERG in Response to X-Rays and to Light

The retina of Rana pipiens, the leopard frog or grass frog, is shown to be an extremely sensitive detector of x-rays. Its sensitivity to x-rays equals in some respects its sensitivity to visible light. The energy required for the response to visible light is so low that the reaction has long been known as one of the most sensitive in biological systems. An exact comparison is made of the amount of energy required in the stimulus to elicit an electroretinogram (ERG) in response to x-rays and in response to light. ERG's from threshold responses to maximal responses obtainable with x-rays and with light are reproduced. The rods of the retina are shown to be responsible for the production of the ERG. The actual amount of energy absorbed in the rhodopsin from x-ray and from light stimulation over a wide range of intensities and durations has been determined and has been related to the amplitude of the ERG. To the question whether light or x-rays are more efficient in eliciting an ERG, no simple or unequivocal answer can be given. The three dimensional relationship of amplitude of response, intensity of stimulus, and duration of stimulus shows rather unexpectedly that in certain regions light is more efficient while in other regions x-rays are more efficient.     

An interplexiform cell in the goldfish retina: light-evked response pattern and intracellular staining with horseradish peroxidase

Summary The light-evoked response pattern and morphology of one interplexiform cell were studied in the goldfish retina by intracellular recording and staining. The membrane potential of the cell spontaneously oscillated in the dark. In response to a brief light stimulus, the membrane potential initially gave a slow transient depolarization. During maintained light, the oscillations showed a tendency to be suppressed; the response of the cell to the offset of the stimulus was not so prominent. The perikaryon of the interplexiform cell was positioned at the proximal boundary of the inner nuclear layer. The cell had two broad layers of dendrites; one was diffuse in the inner plexiform layer, the other was more sparse in the outer plexiform layer. The morphological and electrophysiological characteristics of the cell are discussed in relation to dopaminergic interplexiform cells and the light-evoked release pattern of dopamine in the teleost retina.     

The Rat Electroretinogram : I. Contrasting effects of adaptation on the amplitude and latency of the b-wave

Characteristics of the electroretinogram (ERG) produced by the essentially all rod eye of the rat are presented as functions of the number of quanta absorbed by each rod per stimulus flash. The ERG's were obtained with 1.5 msec. stimulus flashes and uniform illumination of the entire retina. Under these conditions, distortions in the ERG due to stray light are minimized, and the ERG more accurately reflects the activity of its retinal sources. The effects of background light and two forms of dark adaptation were studied and compared. The results, especially for the b-wave, permit an interpretation in terms of two distinct processes. One process appears to determine the b-wave latency. This process is almost independent of the state of adaptation of the retina. The other process does not affect the latency, but determines the b-wave threshold and amplitude. This process strongly depends upon the state of adaptation. Moreover, the effects of dark adaptation on this amplitude-determining process are almost identical with the effects of background light.     

Opponent and nonopponent contributions to the zebrafish electroretinogram using heterochromatic flicker photometry

While some lower vertebrates, such as zebrafish, do not appear to possess anatomically separate pathways of processing visual information (such as M-pathways and P-pathways), it is believed that separate processing of the visual stimulus (such as luminance and chromatic processing) is a basic requirement of vertebrate vision. In this study, spectral sensitivity functions were obtained from electroretinogram responses to heterochromatic flicker photometry stimuli at several flicker rates, including a low flicker rate (2 Hz), in an attempt to predominantly stimulate chromatic processes and a high flicker rate (16 Hz), in an attempt to predominantly stimulate luminance processes. In addition, chromatic adaptation was used to isolate and examine the temporal properties of the different cone-type contributions to the electroretinogram response. Spectral sensitivity functions based on responses to heterochromatic stimuli of a low flicker rate appeared to receive both opponent and nonopponent contributions; however, when the stimulus flicker rate was high, spectral sensitivity appeared to be a function of only nonopponent mechanisms. Also, the differences in cone contributions to the spectral sensitivity functions across the different flicker rates appear to be related to the temporal properties of the cone contributions to the electroretinogram response.     

Serotonin-containing neurones in vertebrate retinas

Abstract: It has been established by a combination of HPLC and electrochemical detection that frog, lizard, goldfish, rabbit, and bovine retinas contain both dopamine and serotonin. Immunohistological and immunoradiographical methods show that serotonin is localised in amacrine perikarya and processes situated in the inner plexiform layers of frog, lizard, and goldfish retinas. The amount of serotonin in the mammalian retina appears to be too low for detection in neurones. The serotonin in the bovine retina is located mainly in the inner nuclear and plexiform layers, suggesting that the amine is present in the same types of cells as found for frog, lizard, and goldfish retinas. Retinas incubated in [³H]serotonin showed that radioactivity is associated with processes in the inner plexiform layer and amacrine perikarya. These results suggest that the neuronal elements that contain endogenous serotonin also have the capacity to accumulate exogenous amine and are consistent with the opinion that serotonin has a neuronal function in retinas of a variety of vertebrates.     

The Spatial Properties of L- and M-Cone Inputs to Electroretinograms That Reflect Different Types of Post-Receptoral Processing

We studied the spatial arrangement of L- and M-cone driven electroretinograms (ERGs) reflecting the activity of magno- and parvocellular pathways. L- and M-cone isolating sine wave stimuli were created with a four primary LED stimulator using triple silent substitution paradigms. Temporal frequencies were 8 and 12 Hz, to reflect cone opponent activity, and 30, 36 and 48 Hz to reflect luminance activity. The responses were measured for full-field stimuli and for different circular and annular stimuli. The ERG data confirm the presence of two different mechanisms at intermediate and high temporal frequencies. The responses measured at high temporal frequencies strongly depended upon spatial stimulus configuration. In the full-field conditions, the L-cone driven responses were substantially larger than the full-field M-cone driven responses and also than the L-cone driven responses with smaller stimuli. The M-cone driven responses at full-field and with 70° diameter stimuli displayed similar amplitudes. The L- and M-cone driven responses measured at 8 and 12 Hz were of similar amplitude and approximately in counter-phase. The amplitudes were constant for most stimulus configurations. The results indicate that, when the ERG reflects luminance activity, it is positively correlated with stimulus size. Beyond 35° retinal eccentricity, the retina mainly contains L-cones. Small stimuli are sufficient to obtain maximal ERGs at low temporal frequencies where the ERGs are also sensitive to cone-opponent processing.     

Intrinsic cone adaptation modulates feedback efficiency from horizontal cells to cones.

Processing of visual stimuli by the retina changes strongly during light/dark adaptation. These changes are due to both local photoreceptor-based processes and to changes in the retinal network. The feedback pathway from horizontal cells to cones is known to be one of the pathways that is modulated strongly during adaptation. Although this phenomenon is well described, the mechanism for this change is poorly characterized. The aim of this paper is to describe the mechanism for the increase in efficiency of the feedback synapse from horizontal cells to cones. We show that a train of flashes can increase the feedback response from the horizontal cells, as measured in the cones, up to threefold. This process has a time constant of approximately 3 s and can be attributed to processes intrinsic to the cones. It does not require dopamine, is not the result of changes in the kinetics of the cone light response and is not due to changes in horizontal cells themselves. During a flash train, cones adapt to the mean light intensity, resulting in a slight (4 mV) depolarization of the cones. The time constant of this depolarization is approximately 3 s. We will show that at this depolarized membrane potential, a light-induced change of the cone membrane potential induces a larger change in the calcium current than in the unadapted condition. Furthermore, we will show that negative feedback from horizontal cells to cones can modulate the calcium current more efficiently at this depolarized cone membrane potential. The change in horizontal cell response properties during the train of flashes can be fully attributed to these changes in the synaptic efficiency. Since feedback has major consequences for the dynamic, spatial, and spectral processing, the described mechanism might be very important to optimize the retina for ambient light conditions.     

The spectral sensitivity of eye movements in response to light flashes inDaphnia magna

Summary The crustaceanDaphnia magna responds to a flash of light with a ventral rotation of its compound eye; this behavior is termed eye flick. We determined the spectral sensitivity for the threshold of eye flick in response to light flashes having three different spatial characteristics: (1) full-field, extending 180° from dorsal to ventral in the animal's field of view; (2) dorsal, 30° wide and located in the dorsal region of the visual field; (3) ventral, same as dorsal but located ventrally. All three stimuli extended 30° to the right and to the left of the animal's midplane. We found that spectral sensitivity varies with the spatial characteristics of the stimulus. For full-field illumination, the relative sensitivity was maximal at 527 nm and between 365 nm and 400 nm, with a significant local minimum at 420 nm. For the dorsal stimulus, the relative sensitivity was greatest at 400 nm, but also showed local maxima at 440 nm and 517 nm. For the ventral stimulus, the relative sensitivity maxima occurred at the same wavelengths as those for the full-field stimulus. At wavelengths of 570 nm and longer, the responses to both dorsal and ventral stimuli showed lower relative sensitivity than the full-field stimulus. No circadian or other periodic changes in threshold spectral sensitivity were observed under our experimental conditions. Animals which had their nauplius eyes removed by means of laser microsurgery had the same spectral sensitivity to full-field illumination as normal animals. Our results are discussed in terms of our current knowledge of the spectral classes of photoreceptors found in theDaphnia compound eye.     

Microscopic investigation of the comparative morphology of the retina

Morphological differences in the architectonics (the relations and composition of the layers and sublayers) of the retina are described in various vertebrates: pike, frog, and cat. These differences apply to both cellular and plexiform layers. The differences are particularly marked in the composition of the sublayers of the inner nuclear layer. In the frog the greatest degree of subdivision into layers of processes of the ganglion and amacrine cells is observed to correspond to the particularly complex differentiation of the inner plexiform layer of the retina (about 10 sublayers). In all the animals studied the ganglion cells can be divided into two principal types: symmetrical and asymmetrical, with many varieties. Asymmetrical amacrine cells are found in the pike and frog retina. The presence of vertical processes branching in the outer plexiform layer is confirmed for amacrine cells in the cat retina. The structural features of the retina are discussed in connection with physiological findings.     

Light-induced changes of sensitivity in Limulus ventral photoreceptors

The responses of Limulus ventral photoreceptors to brief test flashes and to longer adapting lights were measured under voltage clamp conditions. When the cell was dark adapted, there was a range of energy of the test flashes over which the peak amplitude of the responses (light-induced currents) was directly proportional to the flash energy. This was also true when test flashes were superposed on adapting stimuli but the proportionality constant (termed peak currently/photon) was reduced. The peak current/photon was attenuated more by brighter adapting stimuli than by less bright adapting stimuli. The peak current/photon is a measure of the sensitivity of the conductance-increase mechanism underlying the light response of the photo-receptor. The response elicited by an adapting stimulus had a large initial transient which declined to a smaller plateau. The peak current/photon decreased sharply during the declining phase of the transient and was relatively stable during the plateau. This indicates that the onset of light adaptation is delayed with respect to the onset of the response to the adapting stimulus. If the adaptational state just before the onset of each of a series of adapting stimuli was constant, the amplitude of the transient was a nearly linear function of intensity. When the total intensity was rapidly doubled (or halved) during a plateau response, the total current approximately doubled (or halved). We argue that the transition from transient to plateau, light-elicited changes of threshold, and the nonlinear function relating the plateau response to stimulus intensity all reflect changes of the responsiveness of the conductance-increase mechanism.     

The response to monochromatic light flashes of the oxygen consumption of honeybee drone photoreceptors

Local measurements of the fall in oxygen pressure on stimulation of slices of the retina of the honeybee drone by flashes of light were made with oxygen microelectrodes and used to calculate the kinetics of the extra oxygen consumption (delta QO2) induced by each flash. The action spectrum for delta QO2 was obtained from response-intensity curves in response to brief (40 ms) monochromatic light flashes. The action spectrum of receptor potentials was obtained with the same experimental conditions. The two action spectra match closely: they deviate slightly from the photosensitivity spectrum of the drone rhodopsin (R). The deviation is thought to be due to wavelength-dependent light scattering and absorption in the preparation. In these experiments, the visual pigment was first illuminated with orange light, which is known to convert the bistable drone photopigment predominantly to the R state from the metarhodopsin (M) state. When long (300-900 ms) light flashes were used to elicit delta QO2, the responses to different wavelengths could not be matched in time course (as for the short flashes). Flashes producing large R-to-M conversions produced a prolonged delta QO2. The prolongation did not occur after double flashes, which produced both large R-to-M and M-to-R conversions. Similar changes in the length of afterpotentials in the photoreceptor cells and in a long-lasting decrease in photoreceptor intracellular K+ activity were found after long single or double flashes. The results are interpreted to show that the initial event for stimulation by light of metabolism in the drone retina is the same as that for stimulation of electrical responses (i.e., absorption of photons by R). Absorption of photons by M can produce an inhibitory effect on this stimulation.     

Adaptation to Changes in Higher-Order Stimulus Statistics in the Salamander Retina

Adaptation in the retina is thought to optimize the encoding of natural light signals into sequences of spikes sent to the brain. While adaptive changes in retinal processing to the variations of the mean luminance level and second-order stimulus statistics have been documented before, no such measurements have been performed when higher-order moments of the light distribution change. We therefore measured the ganglion cell responses in the tiger salamander retina to controlled changes in the second (contrast), third (skew) and fourth (kurtosis) moments of the light intensity distribution of spatially uniform temporally independent stimuli. The skew and kurtosis of the stimuli were chosen to cover the range observed in natural scenes. We quantified adaptation in ganglion cells by studying linear-nonlinear models that capture well the retinal encoding properties across all stimuli. We found that the encoding properties of retinal ganglion cells change only marginally when higher-order statistics change, compared to the changes observed in response to the variation in contrast. By analyzing optimal coding in LN-type models, we showed that neurons can maintain a high information rate without large dynamic adaptation to changes in skew or kurtosis. This is because, for uncorrelated stimuli, spatio-temporal summation within the receptive field averages away non-gaussian aspects of the light intensity distribution.     

Differences in functional organization of the visual center in frogs and cats

Functional characteristics of responses of tectal neurons in the frog and the primary visual cortex of the cat obtained under indentical experimental conditions during changes in the brightness and duration of flashes within the range of 6 log units were compared. In frogs most units have short summation times and relatively lower thresholds; the response latency is 5–7 times longer than in cats. The overwhelming majority of tectal units can respond only to a narrow range of photic energy that differs for different cells. Most cells in cats respond to changes in brightness of between 4 and 5 log units; they have long summation times, short latent periods, and relatively higher thresholds. The differences found on comparison of the various functional characteristics of the cells in the visual center show that frogs have fixed mechanisms of temporal and spatial interaction responsibile for detection of stimulus brightness. In cats this interaction between individual cell populations and this mutual inhibition between adjacent cells are not prominent. The increased complexity and overlapping of interneuronal connections leads to convergence of day and twilight vision stimuli on the same neuron and to the ability of single units to respond to the whole "working" range of light brightness for the cat visual system.     

Dynamics of turtle horizontal cell response

The small- and large-field (cone) horizontal cells produce similar dynamic responses to a stimulus whose mean luminance is modulated by a white-noise signal. Nonlinear components increase with an increase in the mean luminance and may produce a mean square error (MSE) of up to 15%. Increases in the mean luminance of the field stimulus bring about three major changes: the incremental sensitivity defined by the amplitude of the kernels decreases in a Weber-Fechner fashion; the waveforms of the kernels are transformed from monophasic (integrating) to biphasic (differentiating); the peak response time of the kernels becomes shorter and the cells respond to much higher-frequency inputs. The dynamics of the horizontal cell response also depend on the area of the retina stimulated. Smaller spots of light produce monophasic kernels of a longer peak response time. The presence of a steady background produces three major changes in the spot kernels: the kernel's amplitude becomes larger (incremental sensitivity increases); the peak response times become shorter; the waveform of the kernels changes in a fashion similar to that observed with an increase in the mean luminance of the field stimulus. A similar enhancement in the incremental sensitivity by a steady background has also been observed in catfish, which shows that this phenomenon is a common feature of the horizontal cells in the lower vertebrate retina.     

The functional connections of the supraoptic and suprachiasmatic nuclei of the anterior hypothalamus with the superior colliculi

Interrelation of single hypothalamic (supraoptic--SO and suprachiasmatic--SCH nuclei) and light stimuli at the level of superior colliculus (SC) were studied during chronic experiments in waking rabbits. Short-term inhibitory and the subsequent facilitatory effects of the hypothalamic stimuli on SC responses induced by light flashes were established. The foregoing light stimulus caused inhibitory effect on formation of hypothalamo-collicular responses induced by SO and SCH nuclei stimulation. The final result of the anterior hypothalamic effect on SC function is the vector value of the mechanism of interrelation between the hypothalamic inputs in this structure.