Effect of External Calcium Concentration on the Intensity Dependence of Light-Induced Membrane Current and Voltage Signals in Two Defined States of Adaptation in the Photo-Receptor of Limulus

1986 ◽  
Vol 41 (11-12) ◽  
pp. 1092-1110 ◽  
Author(s):  
H. Stieve ◽  
H. Gaube ◽  
J. Klomfaß

Abstract The intensity dependence of the response of the Limulus ventral nerve photoreceptor to light flashes was determined in alternating measurements for the membrane current signal (receptor current) under voltage clamp conditions and the membrane voltage signal (receptor potential). Responses were obtained at two reproducible states of adaptation, while the photoreceptor was superfused by physiological saline (10 mmol/l Ca2+), or by salines with either lowered (250 μmol/l) or raised (40 mmol/l) calcium concentration. For the dark-adapted state of the photoreceptor the double logarithmic plot of the response current-time integral F (or the current amplitude) vs. flash intensity rises in a steep, supralinear section (slope 2-4) to a curve knee towards a less steep, sublinear section (slope 0.2-0.6), but does not reach saturation in the intensity range tested. Light adaptation shifts the response size vs. intensity curve towards higher light intensities. This sensitivity shift is enlarged in raised, and almost abolished in low external [Ca2+]. The changes of response latency and time-to-peak with stimulus intensity or adaptation are almost identical for receptor current and receptor potential. The decrease-time of the receptor current response, however, depends much less on the stimulus intensity or the state of adaptation than that of the receptor potential. The relative changes in the time course of the receptor current caused by light adaptation are not much influenced by variation of the [Ca2+]ex. Interpretation: The macroscopic receptor current signal consists of a volley of overlapping bumps; the size of these bumps is scaled by a calcium-dependent attenuation function which increases with delay time. This gradual growing attenuation a(t) acts as automatic gain control and may be responsible for the sublinear slope of the intensity dependence of the size of the receptor current. The supralinear slope of this dependence at lower stimulus intensities is probably caused by cooperative effects. Changes in the time course of the macroscopic receptor current due to light adaptation or varied calcium concentration are based on changes in the latency distribution of the underlying bump volley, and the size of the attenuation function.

1980 ◽  
Vol 35 (3-4) ◽  
pp. 308-318 ◽  
Author(s):  
H. Stieve ◽  
I. Claßen-Linke

Abstract The electroretinogram (ERG) of the isolated retina of the crayfish Astacus leptodactylus evoked by strong 10 ms light flashes at constant 5 min intervals was measured while the retina was continuously superfused with various salines which differed in Ca2+ -and Na+ -concentrations. The osmotic pressure of test- and reference-saline was adjusted to be identical by adding sucrose. Results: 1. Upon raising the calcium-concentration of the superfusate in the range of 20-150 mmol/l (constant Na+ -concentration: 208 mmol/l) the peak amplitude hmax and the half time of decay t2 of the ERG both decrease gradually up to about 50% in respect to the corresponding value in reference saline. 2. The recovery of the ERG due to dark adaptation following the “weakly light adapted state” is greatly diminished in high external [Ca2+]ex. 3. Lowering the external calcium-concentration (10 →1 mmol/l) causes a small increase in hmax and a strong increase of the half time of decay t2 (about 180%). Upon lowering the calcium concentration of the superfusate to about 1 nmol/l by 1 mmol/l of the calcium buffer EDTA, a slowly augmenting diminution of the ERG height hm SLX occurs. How­ever, a strong retardation of the falling phase of the ERG characterized by an increase in t2 occurs quickly. Even after 90 min stay in the low calcium saline the retina is still not inexcitable; hmax is 5 - 10% of the reference value. The diminution of hmax occurs about six-fold faster when the buffer concentration is raised to 10 mmol/l EDTA. 4. Additional lowering of the Na+ -concentration (208 →20.8 mmol/l) in a superfusate with a calcium concentration raised to 150 mmol/l causes a strong reduction of the ERG amplitude hmax to about 10%. 5. In a superfusate containing 1 nmol/l calcium such lowering of the sodium concentration (208 → 20.8 mmol/l) causes a diminution of the ERG height to about 40% and the shape of the ERG to become polyphasic; at least two maxima with different time to peak values are observed. Interpretation: 1. The similarity of effects, namely raising external calcium concentration and light adaptation on the one hand and lowering external calcium and dark adaptation on the other hand may indicate that the external calcium is acting on the adaptation mechanism of the photoreceptor cells, presumably by influencing the intracellular [Ca2+]. 2. The great tolerance of the retina against Ca2+ -deficiency in the superfusate might be effected by calcium stores in the retina which need high Ca2+ -buffer concentrations in the superfusate to become exhausted. 3. In contrast to the Limulus ventral nerve photoreceptor there does not seem to be an antagonis­ tic effect of sodium and calcium in the crayfish retina on the control of the light channels. 4. The crayfish receptor potential seems to be composed of at least two different processes. Lowering calcium-and lowering external sodium-concentration both diminish the height and change the time course of the two components to a different degree. This could be caused by in­ fluencing the state of adaptation and thereby making the two maxima separately visible.


1996 ◽  
Vol 107 (6) ◽  
pp. 715-730 ◽  
Author(s):  
M D Gomez ◽  
E Nasi

The receptor potential of rhabdomeric photoreceptors is mediated primarily by a Na influx, but other ions must also permeate through light-dependent channels to account for some properties of the photoresponse. We examined ion conduction in macroscopic and single-channel light-induced currents of slug and scallop photoreceptors. In the absence of Na, a fivefold change in extracellular K shifted the reversal voltage of the photocurrent (Vrev) by approximately 27 mV. Because the dependency of Vrev on [K]o was sub-Nernstian, and Vrev in each condition was more positive than Ek, some other ion(s) with a positive equilibrium potential must be implicated, in addition to K. We assessed the participation of calcium, an important candidate because of its involvement in light adaptation. Three strategies were adopted to minimize the impairments to cytosolic Ca homeostasis and loss of responsiveness that normally result from the required ionic manipulations: (a) Internal dialysis with Na-free solutions, to prevent reverse operation of the Na/Ca exchanger. (b) Rapid solution changes, temporally limiting exposure to potentially detrimental ionic conditions. (c) Single-channel recording, exposing only the cell-attached patch of membrane to the test solutions. An inward whole-cell photocurrent could be measured with Ca as the only extracellular charge carrier. Decreasing the [Ca]o to 0.5 mM reduced the response by 43% and displaced the reversal potential by -4.3 mV; the shift was larger (delta Vrev = -44 mV) when intracellular permeant cations were also removed. In all cases, however, the current carried by Ca was < 5% of that measured with normal [Na]o. Unitary light-activated currents were reduced in a similar way when the pipette contained only divalent cations, indicating a substantial selectivity for Na over Ca. The fall kinetics of the photoresponse was slower when external Ca was replaced by Ba, or when the membrane was depolarized; however, dialysis with 10 mM BAPTA failed to antagonize this effect, suggesting that mechanisms other than the Ca influx participate in the modulation of the time course of the photocurrent.


1966 ◽  
Vol 49 (6) ◽  
pp. 1199-1208 ◽  
Author(s):  
WILLIAM L. PAK ◽  
THOMAS G. EBREY

The second phase (negative peak) of the early receptor potential of cones has been studied in the all-cone eyes of the Mexican and antelope ground squirrels (Citellus mexicanus and Citellus leucurus) and compared with responses from the rod-dominant eyes of the rat and flying squirrel (Glaucomys volans). The responses obtained from the all-cone eyes tended to be smaller in amplitude, to have higher thresholds, and to be considerably more resistant to light adaptation than the responses from the rod-dominant eyes. The wave forms and time courses of the two types of responses were similar, although the cone potential tended to be less sensitive to temperature variations and its time constants tended to be shorter than those of the rod potential. The spectral sensitivity of the second phase of the early receptor potential of the Mexican ground squirrel closely follows the absorption spectrum of a Dartnall nomogram pigment having its absorption maximum at 540 mμ. Moreover, as in the case of the rat, the amplitude of the response appears to be linearly related to the amount of pigment bleached in a flash. Thus, in both all-rod and all-cone systems the early receptor potential appears to arise in the photoexcitation of the respective visual pigment and appears to be closely linked to the initial photochemical events. The similarity of the wave form, time course, and stimulus-response curves in the two systems suggests that the early receptor potential is produced by similar mechanisms in all-rod and all-cone systems.


1999 ◽  
Vol 16 (1) ◽  
pp. 91-105 ◽  
Author(s):  
FRANK NAARENDORP ◽  
GEORGE E. WILLIAMS

We studied the off-response of the rat ERG evoked with long duration, mesopic stimuli during light and dark adaptation, and after intravitreal injection of aspartate and (±)-cis-piperidine-2,3-dicarboxilic acid (PDA). At stimulus offset, the dark-adapted ERG always showed a rapid negative deflection followed by a positive deflection after which the potential returned to baseline. When the stimulus was turned off in the presence of a background of scotopic intensity, the positive deflection consisted of two components. One component was relatively small, fast, and insensitive to rod light adaptation. It resembled the d-wave of the rod ERG. The other component was slow and its amplitude grew with rod light adaptation. In the presence of aspartate, the fast-positive component was absent from the ERG while the remaining positive-going decay of the receptor potential had a time course similar to that of the slow-positive component in the untreated eye. Scotopically matched red and blue stimuli of mesopic intensity elicited equal ERG responses from the dark-adapted eye, including the two positive components in the off-response. These stimuli were also used to assess changes in the ERG off-response during recovery from a strong bleach. Even though the cone contribution to the rat ERG is very small, the presence of a small positive-going component in the off-response following an intense bleach suggested that this response originated from the cone pathway. PDA which suppresses the light response of hyperpolarizing bipolar cells and horizontal cells selectively eliminated the fast-positive component from the ERG. The findings of this study are inconsistent with the idea that the d-wave reflects the decay of the rod receptor potential. They support the possibility that signals from rods cross rod–cone gap junctions at mesopic light intensities, and drive second-order neurons in the cone pathway.


1986 ◽  
Vol 41 (5-6) ◽  
pp. 657-667 ◽  
Author(s):  
I. Claßen-Linke ◽  
H. Stieve

The time course of the recovery of the sensitivity of the Limulus ventral nerve photoreceptor was measured during dark adaptation following light adaptation by a bright 1 or 5 s illumination. The stimulus intensity ICR of a 300 μs light flash evoking a response of criterion amplitude (receptor potential or receptor current under voltage clamp conditions) was used as measure of sensitivity.The time course of dark adaptation shows two phases with time constants in the range of 5-9 s and 300-500 s (15 °C). Only the first of the two phases is significantly changed when the extracel- lular Ca2+-concentration is varied.The power function ICR = a·Io-tDA-b gives a good data fit for each of the two phases of dark adaptation. In the first phase the factor ax and the exponent bx are decreased when the external calcium is lowered from 10 mmol/1 to 250 μmol/1. Conversely a1 and b1 are increased when the Ca2+-concentration is raised to 40 mmol/1. For the second phase neither a2 nor b2 is changed significantly upon the changes in calcium concentration in the same experiments.The two phases of dark adaptation reflect the behaviour of the two components C1 and C2 of the electrical light response (receptor potential or receptor current). Under the conditions described here C, determines the size of the light response during the first phase of dark adaptation whereas C2 mainly influences the size of the response during the second phase.Interpretation: The fast first phase of dark adaptation is determined by the change in intracellu- lar Ca2+-concentration. The slower second phase of dark adaptation is not primarily calcium- controlled.


1983 ◽  
Vol 38 (5-6) ◽  
pp. 471-483 ◽  
Author(s):  
H. Stieve ◽  
H. Gaube

1.Lowering the extracellular calcium concentration from 10 mmol/l to 1 nmol/l causes, besides reducing membrane potential (PMP) and peak amplitude (hmax) of the light response of the Limulus ventral nerve photoreceptor (see Stieve and Bruns [1]), a prolongation of the time course of the light response. The retarded time course (characterized by latent-period tlat, time-to-peak tmax, decrease time t2 and decline quotient QHN) caused by low calcium concentration is not antagonized by either reducing the sodium concentration (from 0.5 to 0.05 mol/l) or increasing the magnesium concentration (from 5× 10-5 to 5 × 10-2 mol/l) in contrast to the effects on the PMP and hmax. 2.This effect of lowering the calcium concentration on the time course of the ReP is distinctly different from that on membrane potential and transient of the ReP described before. It is not characterized by a calcium/sodium binding competition but is probably more closely related to the bump-generating mechanism. It can be explained on the assumption that the time parameters of the ReP are primarily determined by the latency distribution of the underlying bumps which is expanded under low calcium conditions.


1977 ◽  
Vol 66 (1) ◽  
pp. 1-14
Author(s):  
K. Brandle

1. Artifically metamorphosed axolotls were exposed to both brief (impulse) and long-lasting horizontal angular accelerations on a turn-table. The animals responded with a head-turning reaction. 2. The general course of the reaction to impulse acceleration was independent of stimulus intensity. The velocity of the head movement first increased to a maximum exponentially and then decreased in a negative exponential manner. Stimulus intensity had a linear relationship to the mean maximum velocity and mean total angle covered by head-turning. The average velocity-time curves at various stimulus intensities differed only by a velocity factor. 3. During long-lasting constant accelerations the velocity of the head-turning increased to a maximum velocity in a sigmoid time-course and then decreased, first to a constant velocity, and then further. Mean values of the maximum velocity were correlated linearly with the stimulus intensity. 4. It was concluded that the head-turning reflexes in axolotls do not agree with the accepted movements of the vertebrate cupula and therefore are not a simple ‘copy’ of the afferent input. It is also suggested that the reaction threshold differes from that for the labyrinthine input.


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