Light adaptation and color opponency of horizontal cells in the turtle retina

Chromaticity-type (C-type) horizontal cells of the turtle retina receive antagonistic inputs from cones of different spectral types, and therefore their response to background illumination is expected to reflect light adaptation of the cones and the interactions between their antagonistic inputs. Our goal was to study the behavior of C-type horizontal cells during background illumination and to evaluate the role of wavelength in background adaptation. The photoresponses of C-type horizontal cells were recorded intracellularly in the everted eyecup preparation of the turtleMauremys caspicaduring chromatic background illuminations. The voltage range of operation was either reduced or augmented, depending upon the wavelengths of the background and of the light stimuli, while the sensitivity to light was decreased by any background. The response–intensity curves were shifted to brighter intensities and became steeper as the background lights were made brighter regardless of wavelength. Comparing the effects of cone iso-luminant backgrounds on the Red/Green C-type horizontal cells indicated that background desensitization in these cells could not solely reflect background adaptation of cones but also depend upon response compression/expansion and changes in synaptic transmission. This leads to wavelength dependency of background adaptation in C-type horizontal cells, that is expressed as increased light sensitivity (smaller threshold elevation) and improved suprathreshold contrast detection when the wavelengths of the background and light stimuli were chosen to exert opponent effects on membrane potential.

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Turtle C-type horizontal cells act as push–pull devices

Visual Neuroscience ◽

10.1017/s0952523801186050 ◽

2001 ◽

Vol 18 (6) ◽

pp. 893-900 ◽

Cited By ~ 3

Author(s):

G. TWIG ◽

H. LEVY ◽

I. PERLMAN

Keyword(s):

Short Wavelength ◽

Horizontal Cells ◽

Background Illumination ◽

Voltage Range ◽

Long Wavelength ◽

Relative Contribution ◽

Color Opponency ◽

Light Stimuli ◽

Short Wavelength Light ◽

Wavelength Light

Chromaticity (C-type) horizontal cells in the retina of cold-blooded vertebrates receive antagonistic inputs from cone photoreceptors of different spectral types leading to color opponency. The relative contribution of each spectral type of cones can be selectively altered by chromatic background illumination. Therefore, the spectral properties of C-type horizontal cells are expected to change when the intensity and color of ambient illumination are altered. In this study, we investigated the effects of chromatic background lights upon color opponency in Red/Green (RGH) and Yellow/Blue (YBH) C-type horizontal cells in the everted eyecup preparation of the turtle Mauremys caspica. Photoresponses were elicited by long-wavelength and short-wavelength light stimuli in the dark-adapted state and under conditions of chromatic background illumination. We found that the total voltage range, within which graded depolarizing and the hyperpolarizing photoresponses could be elicited, either increased or decreased depending upon the color of the background light. However, the maximal and minimal potential levels determined respectively by long-wavelength and short-wavelength light stimuli of supersaturating intensity remained unchanged, regardless of the wavelength and intensity of the background. These findings indicate that turtle C-type horizontal cells operate as push–pull devices. A sufficiently bright short-wavelength stimulus can push them all the way to the maximal hyperpolarizing level while a very bright long-wavelength stimulus can pull them towards the most depolarizing potential.

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Effects of light adaptation on contrast processing in bipolar cells in the retina

Visual Neuroscience ◽

10.1017/s0952523801184087 ◽

2001 ◽

Vol 18 (4) ◽

pp. 581-597 ◽

Cited By ~ 9

Author(s):

PATRICK K. FAHEY ◽

DWIGHT A. BURKHARDT

Keyword(s):

Light Adaptation ◽

Dynamic Range ◽

Horizontal Cell ◽

Bipolar Cells ◽

Horizontal Cells ◽

Background Intensity ◽

Ambystoma Tigrinum ◽

Background Illumination ◽

Voltage Range ◽

Contrast Gain

Effects of light adaptation on contrast processing in the outer retina were investigated over nearly four decades of background illumination by analyzing the intracellular responses of 111 bipolar cells, 66 horizontal cells, and 22 cone photoreceptors in the superfused eyecup of the tiger salamander (Ambystoma tigrinum). Light adaptation had striking and similar effects on the average contrast responses of the hyperpolarizing (Bh) and depolarizing (Bd) classes of bipolar cells: Over the lower two decades of background illumination, the contrast gain increased 7-fold to reach values as high as 20–30, the dynamic range and the half-maximum contrast decreased by about 60%, the total voltage range increased some 40%, and contrast dominance changed from highly positive to more balanced. At higher levels of background, most aspects of the contrast response stabilized and Weber's Law then held closely. In this background range, the contrast gain of bipolar cells was amplified some 20× relative to that of cones whereas the corresponding amplification in horizontal cells was about 6×. Differences in the growth of contrast gain with the intensity of the background illumination for cones versus bipolar cells suggest that there are at least two adaptation-dependent mechanisms regulating contrast gain. One is evident in the cone photoresponse such that an approximately linear relation holds between the steady-state hyperpolarization and contrast gain. The other arises between the voltage responses of the cones and bipolar cells. It could be presynaptic (modulation of cone transmitter release by horizontal cell feedback or other mechanisms) and/or postsynaptic, that is, intrinsic to bipolar cells. Contrast gain grew with the background intensity by a larger factor in horizontal than in bipolar cells. This provides a basis for the widely held view that light adaptation increases the strength of surround antagonism in bipolar cells. On average, the effects of light adaptation and most quantitative indices of contrast processing were remarkably similar for Bd and Bh cells, implying that both classes of bipolar cells, despite possible differences in underlying mechanisms, are about equally capable of encoding all primary aspects of contrast at all levels of light adaptation.

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The effects of GABA and related drugs on horizontal cells in the isolated turtle retina

Visual Neuroscience ◽

10.1017/s0952523800000596 ◽

1990 ◽

Vol 5 (5) ◽

pp. 469-477 ◽

Cited By ~ 19

Author(s):

Ido Perlman ◽

Richard A. Normann

Keyword(s):

Plexiform Layer ◽

Neural Mechanism ◽

Competitive Inhibitor ◽

Horizontal Cells ◽

Gaba Uptake ◽

Uptake System ◽

Cobalt Ions ◽

Color Opponency ◽

Chromatic Properties

AbstractThe role of GABA in the outer plexiform layer of the turtle retina has been examined by intracellular recordings from L- and C-type horizontal cells in the isolated retina preparation.GABA (1–5 mM) slightly depolarized the L-type horizontal cells, reduced the amplitude of their photoresponses, and slowed down the rate of hyperpolarization during the ON component of the photoresponse. These effects could not be replicated by either muscimol or baclofen. When synaptic transmission from the photoreceptors had been blocked by either kynurenic acid or cobalt ions, GABA depolarized L-type horizontal cells and augmented the remaining photoresponses. Neither muscimol nor baclofen exerted any effect on L-type horizontal cells under these conditions. Nipecotic acid, a competitive inhibitor of the GABA-uptake system, induced effects on turtle L-type horizontal cells which were similar to those exerted by GABA. Thus, the complex GABA effect on turtle L-type horizontal cells seems to represent the summation of at least two actions; an indirect one mediated by the red cones via GABAa-type receptors and a direct one which probably reflects the activation of an electrogenic GABA-uptake system.GABA (1–5 mM) induced a transient depolarization in C-type horizontal cells but eliminated color opponency in only three cells out of seven studied. This observation is inconsistent with the notion that the only neural mechanism responsible for the chromatic properties of C-type horizontal cells in the turtle retina is a GABAergic negative feedback from the L-type horizontal cells onto the green ones.

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Sensitivity transformation for vertebrate vision

Visual Neuroscience ◽

10.1017/s0952523800006593 ◽

1991 ◽

Vol 6 (4) ◽

pp. 371-374 ◽

Cited By ~ 4

Author(s):

Richard L. Chappell ◽

Ken-Ichi Naka

Keyword(s):

Light Adaptation ◽

Light Response ◽

Horizontal Cells ◽

Visual Response ◽

Maximum Point ◽

Half Maximum ◽

Peak Response ◽

Vertebrate Retina ◽

Mean Luminance

AbstractThe visual response to a flash given in the dark is known to saturate according to the Michaelis-Menten relationship. Nevertheless, the incremental response from increasing levels of mean luminance tends to follow a Weber-Fechner relationship well into the saturation range determined from the Michaelis-Menten results. This sensitivity transformation from Michaelis-Menten to Weber-Fechner is an important characteristic of light adaptation in the vertebrate retina. Recent studies concerning the role of calcium in photoreceptor adaptation have shown that the relaxation from peak to plateau in the response of isolated photoreceptors was absent under conditions in which adaptation was blocked. Comparing the pronounced relaxation from peak to plateau in turtle horizontal cells with the absence of such relaxation in the catfish response, we noted also that turtle incremental sensitivity shows a Weber-Fechner relationship while catfish incremental sensitivity more closely follows the local slope of the Michaelis-Menten relation. Based on these observations, we have obtained an expression to relate the relaxation from peak to plateau with the sensitivity transformation. We assume that adaptation shifts the half-maximum point of the Michaelis-Menten curve so that the light response relaxes to a plateau value equal to a specified fraction φ of the peak response. We show that this manipulation alone results in a transformation from Michaelis-Menten kinetics to Weber-Fechner sensitivity.

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