Background-induced flicker enhancement in cat retinal horizontal cells. II. Spatial properties

1990 ◽  
Vol 64 (2) ◽  
pp. 326-340 ◽  
Author(s):  
R. Nelson ◽  
R. Pflug ◽  
S. M. Baer
Keyword(s):  
Test Stimulus ◽  
Exponential Decay ◽  
Horizontal Cell ◽  
Stimulus Size ◽  
Slit Width ◽  
Horizontal Cells ◽  
Test Region ◽  
Length Constant ◽  
Region Model ◽  
The Mean

1. Intracellular recordings have been made from cat retinal horizontal cells stimulated with flickering test spots. Dim backgrounds increase flicker amplitudes in response to small but not large test stimuli. 2. This background-induced flicker enhancement has been measured for different slit- and square-test stimulus widths and the results compared with two spatial models for the enhancement effect. 3. In the "dark test-region" model it is argued that rods within the test region are unresponsive to background stimuli because of prior saturation by the test stimulus. Background-evoked rod signals decay passively from regions outside the test stimulus through a syncytial network into the recording site, where they act on the cone-to-horizontal-cell synapse, increasing its gain. 4. In the "changing length-constant" model rod signals reduce the length constant of a syncytial network by uncoupling the cells within it. This causes an increased response to small but not large test stimuli. 5. Both models are analytically evaluated with the use of a conductive-sheet approximation to the syncytial network. Expressions are derived for network polarization [(V(0, 0)] as a function of stimulus size. The specific stimulus shapes considered are disks, rectangles, slits, and squares in both bright and dark varieties. From these expressions predictions of response enhancement as a function of stimulus size are made for both models. 6. The dark test-region model provides for an exponential decay of flicker enhancement as a function of slit width but a steeper-than-exponential decay with the width of squares, in close agreement with experimental data. 7. The changing length-constant model makes qualitatively similar predictions. Flicker enhancement declines nearly exponentially with slit width. For square-shaped test stimuli the predicted decline of flicker enhancement with size is somewhat shallower than either the dark test-region-model curve or the experimentally determined curve. 8. As recorded in the same set of cells and under the same set of stimulus conditions (with the use of both slit- and square-test stimuli), the mean length constant of the peak-to-peak flicker component in the horizontal-cell response is 168 +/- 18 (SE) microns with the background and 232 +/- 45 microns in the dark. The mean length constant for the background-induced flicker enhancement, as fit by dark test-region-model curves, is 186 +/- 22 microns (n = 9).(ABSTRACT TRUNCATED AT 250 WORDS)

Dynamic changes in the receptive fields of L1-type horizontal cells in the retina of the turtle Mauremys caspica

2002 ◽  
Vol 19 (5) ◽  
pp. 621-632 ◽  
Author(s):  
O. BORNSTEIN ◽  
G. TWIG ◽  
J. BENDA ◽  
R. WEILER ◽  
I. PERLMAN
Keyword(s):  
Receptive Field ◽  
Light Stimulus ◽  
Horizontal Cell ◽  
Receptive Fields ◽  
Horizontal Cells ◽  
Time Dependent ◽  
Full Field ◽  
Receptive Field Properties ◽  
Length Constant ◽  
Mauremys Caspica

The resistances of the horizontal cell syncytium in the vertebrate retina are modulated in a time-dependent fashion during light stimulation. Therefore, the spatial properties of horizontal cells are expected to change with time after the illumination conditions are altered. This study was designed to investigate time- and intensity-dependent changes in the receptive-field properties of L1-type horizontal cells in the turtle Mauremys caspica. Photoresponses were elicited by monochromatic (650 nm) light stimuli of 2-s duration covering retinal spots of different radii. The length constants were derived from the relationships between amplitude and spot radius that were constructed for different time intervals after onset of the light stimulus. For a given stimulus intensity, the length constant transiently increased to a peak value and then slowly recovered to a plateau level. When the length constant was compared to the amplitude of the response to full-field illumination for the entire duration of the light stimulus, an ellipse-like curve was obtained indicating that for a given membrane potential, two different values of the length constant could be obtained. Dopamine considerably reduced the size of the receptive fields but did not affect the time-dependent changes in the length constant. These results indicate that changes in the membrane resistance underlie short-term modulation of the receptive-field properties of turtle L1-type horizontal cells after onset of a light stimulus.

scholarly journals Dynamics of turtle horizontal cell response.

1985 ◽  
Vol 86 (3) ◽  
pp. 423-453 ◽  
Author(s):  
R L Chappell ◽  
K Naka ◽  
M Sakuranaga
Keyword(s):  
Response Time ◽  
Cell Response ◽  
Horizontal Cell ◽  
Horizontal Cells ◽  
Dynamic Responses ◽  
Large Field ◽  
Lower Vertebrate ◽  
Peak Response ◽  
The Mean ◽  
Mean Luminance

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.

Modulation of A-type potassium currents in retinal horizontal cells by extracellular calcium and zinc

2006 ◽  
Vol 23 (5) ◽  
pp. 825-832 ◽  
Author(s):  
DAO-QI ZHANG ◽  
ZIYI SUN ◽  
DOUGLAS G. MCMAHON
Keyword(s):  
Steady State ◽  
Potassium Channels ◽  
Horizontal Cell ◽  
Horizontal Cells ◽  
Resting Membrane Potential ◽  
Inactivation Curve ◽  
Outward Currents ◽  
Steady State Inactivation ◽  
The Mean ◽  
Kinetics Of

Extracellular Ca2+ and Zn2+ influence many aspects of retinal function. Here, we examined the effect of external Ca2+ and Zn2+ on potassium channels of retinal horizontal cells. When extracellular Ca2+ was lowered from 3 mM to 0.3 mM, horizontal cell transient outward currents elicited by voltage steps from resting membrane potential (−70 mV) were decreased by approximately 50%, whereas the sustained currents remained unchanged. This effect was due to a hyperpolarizing shift in the steady-state inactivation curve of A-type K+ currents when extracellular Ca2+ concentration was lowered. The mean half inactivation potential of the steady-state inactivation curves was hyperpolarized from −56.3 ± 4.7 mV in 3 mM Ca2+ to −76.4 ± 3.9 mV in 0.3 mM Ca2+. Neither the state-steady activation curve nor the kinetics of inactivation was significantly changed in low extracellular Ca2+. The addition of 30 μM Zn2+ restored peak outward currents in 0.3 mM Ca2+. The half inactivation voltages were depolarized from −70 ± 2.8 mV in 0.3 mM Ca2+ to −56 ± 2.6 mV in 0.3 mM Ca2+ plus 30 μM Zn2+. Taken together, the results indicate that external Ca2+ and Zn2+ maintain the activity of A-type potassium channels in retinal horizontal cells by influencing the voltage dependence of steady-state inactivation.

scholarly journals Lateral feedback from monophasic horizontal cells to cones in carp retina. I. Experiments.

1989 ◽  
Vol 93 (4) ◽  
pp. 681-694 ◽  
Author(s):  
M Kamermans ◽  
B W van Dijk ◽  
H Spekreijse ◽  
R C Zweypfenning
Keyword(s):  
Horizontal Cell ◽  
Horizontal Cells ◽  
Color Coding ◽  
Cell Adaptation ◽  
Test Spot ◽  
Displacement Experiments ◽  
High Stimulus ◽  
Integration Area

The spatial and color coding of the monophasic horizontal cells were studied in light- and dark-adapted retinae. Slit displacement experiments revealed differences in integration area for the different cone inputs of the monophasic horizontal cells. The integration area measured with a 670-nm stimulus was larger than that measured with a 570-nm stimulus. Experiments in which the diameter of the test spot was varied, however, revealed at high stimulus intensities a larger summation area for 520-nm stimuli than for 670-nm stimuli. The reverse was found for low stimulus intensities. To investigate whether these differences were due to interaction between the various cone inputs to the monophasic horizontal cell, adaptation experiments were performed. It was found that the various cone inputs were not independent. Finally, some mechanisms for the spatial and color coding will be discussed.

Receptive field of the retinal bipolar cell: a pharmacological study in the tiger salamander

1996 ◽  
Vol 76 (3) ◽  
pp. 2005-2019 ◽  
Author(s):  
W. A. Hare ◽  
W. G. Owen
Keyword(s):  
Receptive Field ◽  
Gaba Receptors ◽  
Bipolar Cell ◽  
Horizontal Cell ◽  
Pharmacological Study ◽  
Bipolar Cells ◽  
Horizontal Cells ◽  
Gaba Uptake ◽  
Gamma Aminobutyric Acid ◽  
Gabaergic Synapse

1. It is widely believed that signals contributing to the receptive field surrounds of retinal bipolar cells pass from horizontal cells to bipolar cells via GABAergic synapses. To test this notion, we applied gamma-aminobutyric acid (GABA) agonists and antagonists to isolated, perfused retinas of the salamander Ambystoma tigrinum while recording intracellularly from bipolar cells, horizontal cells, and photoreceptors. 2. As we previously reported, administration of the GABA analogue D-aminovaleric acid in concert with picrotoxin did not block horizontal cell responses or the center responses of bipolar cells but blocked the surround responses of both on-center and off-center bipolar cells. 3. Surround responses were not blocked by the GABA, antagonists picrotoxin or bicuculline, the GABAB agonist baclofen or the GABAB antagonist phaclofen, and the GABAC antagonists picrotoxin or cis-4-aminocrotonic acid. Combinations of these drugs were similarly ineffective. 4. GABA itself activated a powerful GABA uptake mechanism in horizontal cells for which nipecotic acid is a competitive agonist. It also activated, both in horizontal cells and bipolar cells, large GABAA conductances that shunted light responses but that could be blocked by picrotoxin or bicuculline. 5. GABA, administered together with picrotoxin to block the shunting effect of GABAA activation, did not eliminate bipolar cell surround responses at concentrations sufficient to saturate the known types of GABA receptors. 6. Surround responses were not blocked by glycine or its antagonist strychnine, or by combinations of drugs designed to eliminate GABAergic and glycinergic pathways simultaneously. 7. Although we cannot fully discount the involvement of a novel GABAergic synapse, the simplest explanation of our findings is that the primary pathway mediating the bipolar cell's surround is neither GABAergic nor glycinergic.

Nitric oxide and cGMP modulate retinal glutamate receptors

1996 ◽  
Vol 76 (4) ◽  
pp. 2307-2315 ◽  
Author(s):  
D. G. McMahon ◽  
L. V. Ponomareva
Keyword(s):  
Nitric Oxide ◽  
Glutamate Receptors ◽  
Horizontal Cell ◽  
Synaptic Function ◽  
Horizontal Cells ◽  
Glutamate Excitotoxicity ◽  
No Donors ◽  
Receptor Modulation ◽  
Cell Recording ◽  
Endogenous Nitric Oxide

1. In the retina, as in other regions of the vertebrate central nervous system, glutamate receptors mediate excitatory chemical synaptic transmission and are a critical site for the regulation of cellular communication. In this study, retinal horizontal cells from the hybrid less were dissociated in cell culture, voltage clamped by the whole cell recording technique, and the currents evoked by application of excitatory amino acids recorded. 2. Responses to glutamate and its agonist kainate were reduced by approximately 50% in the presence of the nitric oxide (NO) donors sodium nitroprusside and S-nitroso-N-acetylpenicillamine. The effect of these compounds was blocked by the NO scavenger hemoglobin. 3. This effect of NO donors on kainate currents could be mimicked by the application of a membrane permeable guanosine 3',5'-cyclic monophosphate (cGMP) analogue, 8-Br-cGMP. The NO effect was also blocked by application of the guanylate cyclase inhibitor LY-83583, and by a protein kinase G inhibitor peptide. 4. In H1-type horizontal cells, stimulation of endogenous nitric oxide synthase with L-arginine reduced kainate responses, whereas application of D-arginine had no effect. 5. This receptor modulation mechanism may act in concert with other pre- and postsynaptic mechanisms to modify horizontal cell synaptic function according to the adaptational state of the retina and also may protect horizontal cells from glutamate excitotoxicity.

scholarly journals Modelling Activity and Durability of Electrocatalysts from a Statistical Perspective

2021 ◽  
Author(s):  
Jun Huang
Keyword(s):  
Exponential Decay ◽  
Analytical Solutions ◽  
Electrocatalytic Activity ◽  
Active Sites ◽  
Simple Theory ◽  
Exact Analytical Solutions ◽  
Modelling Activity ◽  
Closing The Gap ◽  
The Mean ◽  
Kinetics Of

<p>We develop a theory to investigate how energetic nonhomogeneity of active sites determines the overall activity of an electrocatalyst and how the evolution of the nonhomogeneity determines the overall durability. The simple theory is amenable to exact analytical solutions and thus fosters an in-depth transparent analysis. It is revealed that nonhomogeneity does not necessarily diminish the electrocatalytic activity; instead, the highest overall activity is obtained with a suitable level of nonhomogeneity that is commensurate with the mean property. The evolution kinetics of nonhomogeneity is described by using the Fokker-Planck theory. Exponential decay of the activity is predicted theoretically and confirmed experimentally. The present work represents a first step toward closing the gap between model and practical electrocatalysts using statistical considerations.</p>

3H-adenosine uptake selectively labels rod horizontal cells in goldfish retina

1997 ◽  
Vol 14 (2) ◽  
pp. 207-212 ◽  
Author(s):  
Keith M. Studholme ◽  
Stephen Yazulla
Keyword(s):  
Staining Pattern ◽  
Outer Nuclear Layer ◽  
Horizontal Cell ◽  
Horizontal Cells ◽  
The Other ◽  
Axon Terminals ◽  
Adenosine Uptake ◽  
Double Label ◽  
Injection Methods ◽  
Goldfish Retina

AbstractThere are four types of horizontal cell in the goldfish retina, three cone- and one rod-type. The neurotransmitter of only one type, the H1 (cone) horizontal cell, has been identified as GABA. 3H-adenosine uptake was examined as a possible marker for the other classes of horizontal cell. Isolated goldfish retinae were incubated in 3H-adenosine (10–40 μCi) in HEPES-buffered saline for 30 min, then fixed, embedded in plastic, and processed for light-microscopic autoradiography (ARG). For double-label immuno/ARG studies, l-μm-thick sections were processed for GABA postembed immunocytochemistry, then for ARG. 3H-adenosine uptake was localized to cone photoreceptors, presumed precursor cells in the proximal outer nuclear layer, and to a single, continuous row of horizontal cell bodies in the inner nuclear layer. No uptake was localized to the region of horizontal cell axon terminals. 3H-adenosine uptake did not colocalize with GABA-IR in H1 horizontal cells, but it did colocalize with adenosine deaminase immunoreactivity. It is concluded that 3H-adenosine uptake selectively labels rod horizontal cells in the goldfish retina based on position and staining pattern, which are similar to rod horizontal cells stained by Golgi or HRP injection methods. The use of 3H-adenosine uptake may provide a useful tool to study other properties of rod horizontal cells (i.e. development) as well as provide clues as to the transmitter used by these interneurons.

Dual effect of glycine on horizontal cells of the tiger salamander retina

1991 ◽  
Vol 66 (6) ◽  
pp. 1993-2001 ◽  
Author(s):  
S. Borges ◽  
M. Wilson
Keyword(s):  
Opposite Effect ◽  
Horizontal Cell ◽  
Light Response ◽  
Membrane Resistance ◽  
Horizontal Cells ◽  
Field Size ◽  
Dual Effect ◽  
Light Responses ◽  
Low Concentrations ◽  
High Concentrations

1. The effects of glycine on horizontal cells have been examined by microelectrode recording from superfused retinas isolated from the salamander. 2. Low concentrations of glycine (less than 50 microM) hyperpolarized horizontal cells and increased the magnitude of their light responses. Millimolar concentrations produced the opposite effect of depolarizing these cells and reducing their light response amplitudes. 3. In the presence of Co2+ and Mg2+ at concentrations sufficient to suppress the light response, millimolar glycine still exerted a depolarizing effect on horizontal cells, implying that this effect was largely a direct one on horizontal cell membranes. 4. Although both the rod and the cone contributions to horizontal cell light responses were reduced by millimolar glycine, rod input was reduced more, suggesting that millimolar glycine may also exert a presynaptic effect. 5. Strychnine (10 microns) antagonized the effects of millimolar glycine and, in the absence of exogenously applied glycine, caused horizontal cells to hyperpolarize and their light responses to increase in amplitude. This result implies that, in darkness, glycine is tonically released onto horizontal cells and maintains them in a state of partial depolarization. 6. The low-concentration effect of glycine was accompanied by an increased membrane resistance and receptive field size but no change in the balance of rod and cone input. 7. Low concentrations of glycine were often seen to cause a speeding of light responses, whereas high concentrations sometimes caused a slowing of response kinetics. Response kinetics were found to correlate with horizontal cell dark membrane potential so that, positive to -30 mV, depolarization slowed responses whereas kinetics at more negative values were largely independent of voltage.

Bifurcation analysis of nonlinear retinal horizontal cell models. II. Network properties

1990 ◽  
Vol 64 (1) ◽  
pp. 248-261 ◽  
Author(s):  
R. L. Winslow ◽  
S. Ma
Keyword(s):  
Horizontal Cell ◽  
Membrane Currents ◽  
Horizontal Cells ◽  
Synaptic Conductance ◽  
Cell Models ◽  
Voltage Dependent ◽  
Full Complement ◽  
Network Properties ◽  
Light Stimuli ◽  
Coupling Conductance

1. We have previously presented a model of horizontal-cell soma isolated from fish retina. The model consists of a synaptic conductance representing input from photoreceptors in parallel with voltage-dependent membrane currents. Membrane-current models are based on I-V curves measured in isolated fish horizontal cells. Bifurcation theory was used to analyze model properties. The major findings of this study were 1) the inward Ca2+ current must be inactivated to account for horizontal-cell resting potentials and hyperpolarizing responses to light stimuli in a background of dark, and 2) the synaptic conductance controls the bifurcation structure of the model, with bistable behavior occurring at small and monostable behavior occurring at larger values of the synaptic conductance. The synaptic conductance at the point of transition from bistable to monostable behavior corresponds to the activation of as few as 100 synaptic channels. Thus tonic synaptic input from photoreceptors and inactivation of the inward Ca2+ current act to “linearize” responses of isolated horizontal-cell models. 2. The model described in this paper extends these analyses to large networks of horizontal cells in which each cell is coupled resistively to its nearest neighbors and is modeled with the use of the full complement of nonlinear membrane currents. Network responses to arbitrary patterns of conductance change (simulating inputs from photoreceptors), current-, or voltage-clamp stimuli are computed using the Newton iteration. The Newton descent direction is computed using either conjugate gradient (CG) or preconditioned CG algorithms. 3. An analysis of network stability properties is performed. Network I-V curves are computed by voltage-clamping the center node and computing the current required to maintain the clamp voltage. Computations are performed on networks of model cells in which the Ca2+ current is fully activated and the synaptic conductance is zero, thus making each cell as nonlinear as possible. Coupling conductance values slightly greater than 100 pS provide a current shunt sufficient to prevent the generation of Ca2+ action potentials in the network. This coupling conductance corresponds to the conductance of as few as two gap-junction channels and is more than two orders of magnitude less than the coupling known to exist between pairs of cultured horizontal cells.(ABSTRACT TRUNCATED AT 400 WORDS)

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