Search PubMed⌕ Search

Biomedical subjects

V Torre

Publications and source records attributed to V Torre.

At least 73 records · Page 4Linked to original sources

Mechanisms of light adaptation in toad rods.

A steady illumination equivalent to about 1000 or more photoisomerizations per s per rod completely abolishes the dark current and induces a total desensitization of toad rods. Under these conditions, however, both current and sensitivity recover from the initial suppression and reach a new steady level within about 30 s. Both extent and time course of recovery depend on the intensity of the conditioning light. The ability of rods to respond to light stimuli under light saturating conditions is enhanced by procedures aimed at increasing the sodium gradient across their surface membrane. The results suggest that rods can also contribute significantly to vision at photopic levels of ambient light. The mechanisms underlying this adaptational property of rods are discussed.

Acclimatization↗

High-pass filtering of small signals by the rod network in the retina of the toad, Bufo marinus.

The electrical spread of excitation in the network of rod photoreceptors was studied by intracellular recording in the isolated, perfused retina of the toad, Bufo marinus. Experiments with dim, bar-shaped flashes of light revealed that the rod network behaves as a high-pass filter to laterally propagating small signals. Such a behavior had been found earlier in the turtle (Detwiler et al., 1980). Three electrical equivalent circuit models that can explain this behavior were considered and analytical solutions to the network equations were obtained. By fitting these analytical expressions to linear responses elicited by weak light flashes and to voltage excursions elicited by extrinsic current injections, values for the circuit parameters were determined. Values obtained by independent methods were consistent. The effects of changing each of these parameters in turn upon the high-pass filtering of small signals were then predicted. These predictions provided a framework for an analysis of the ionic basis of the underlying mechanism, which is described in the following paper.

Animals↗

High-pass filtering of small signals by retinal rods. Ionic studies.

The high-pass filtering of small signals by the rod photoreceptor network was studied by intracellular recording in the isolated, perfused retina of the toad, Bufo marinus. Data were analyzed and interpreted in terms of the network analysis described in the preceding paper. External concentrations of Cs+ as high as 10 mM, which blocked the relaxation from peak to plateau of the rod's response to bright light, did not affect the filtering of small signals. The effects of reducing [Na+]o were not consistent with a direct action upon the mechanism underlying this filtering property. By contrast, raising external [K+] from 2.6 to 10 mM, which caused a fourfold reduction in EK, abolished the high-pass filtering of small signals. Analysis of the effects of external [K+] changes indicates that the underlying mechanism involves a K+ conductance that decreases with a delay when the rod is hyperpolarized. This conductance is not blocked by externally applied tetraethylammonium. Other experiments did not rule out the possibility that it might be activated by Ca++.

Animals↗

Nonlinear interactions in a dendritic tree: localization, timing, and role in information processing.

In a passive dendritic tree, inhibitory synaptic inputs activating ionic conductances with an equilibrium potential near the resting potential can effectively veto excitatory inputs. Analog interactions of this type can be very powerful if the inputs are appropriately timed and occur at certain locations. We examine with computer simulations the precise conditions required for strong and specific interactions in the case of a delta-like ganglion cell of the cat retina. We find some critical conditions to be that (i) the peak inhibitory conductance changes must be sufficiently large (i.e., approximately equal to 50 nS or more), (ii) inhibition must be on the direct path from the location of excitation to the soma, and (iii) the time course of excitation and inhibition must substantially overlap. Analog AND-NOT operations realized by satisfying these conditions may underlie direction selectivity in ganglion cells.

Action Potentials↗

The effect of phosphodiesterase inhibitors on the electrical activity of toad rods.

The membrane potential of toad rods was recorded during addition of small amounts of phosphodiesterase inhibitors to the extracellular medium. Separate application of 3-isobutyl-1-methylxanthine (IBMX), caffeine, theophylline, papaverine and RO 20-1724 slowed down the time course of rod photo-response to dim flashes of light. These changes were associated with a two to six-fold increase in the amplitude of photoresponse. The effects on kinetics may be described simply by an expansion of the photoresponse time scale. When the drug concentration was raised above a certain level, the rods showed supralinear behaviour whereby doubling of the intensity of a dim flash could increase the response more than two-fold. Under similar conditions rods also showed light sensitization whereby responses to dim flashes were enhanced in the presence of dim backgrounds. Taking the drug concentration that induced a two-fold increase in the time-to-peak, IBMX was found the most effective compound, followed by papaverine, RO 20-1724, theophylline and caffeine with relative effectivities 1, 1/2, 1/7, 1/40 and 1/100. Sensitivity, kinetics and supralinear behaviour may be restored to normal by steady background illumination while still in the presence of IBMX. However the intensity of the steady light, needed to restore the sensitivity to control levels, is not sufficient to accelerate the kinetics back to control values. In the presence of 50 microM-IBMX a dim steady background of light enhanced the response to dim flashes. When the intensity of the light background was increased rods were desensitized and the supralinear behaviour disappeared. The antagonism between the effects of IBMX and the effects of background illumination on the kinetics of photoresponse suggests that phosphodiesterase activity controls the time course of light response in vertebrate rods.

1-Methyl-3-isobutylxanthine↗

Ionic movements through light-sensitive channels of toad rods.

Electrical photoresponses of rods in the isolated toad retina were recorded during ionic manipulations of the Na+-free extracellular medium. In the presence of a concentration of external Ca2+ above 10(-5) M, voltage photoresponses were observed only in the presence of external Na+ or Li+. When external Ca2+ was reduced below 10(-6) M, voltage photoresponses of normal polarity could be detected even in the absence of Na+ or Li+, but in the presence of external Mg2+. In the presence of normal extracellular Ca2+ hyperpolarizing photoresponses were observed even in the absence of Na+ or Li+, provided small amounts of phosphodiesterase inhibitors (IBMX, RO 20-1724, papaverine, caffeine, theophylline) were added to the perfusate. Responses obtained in low-Na+ IBMX solutions required the presence of millimolar amounts of a variety of divalent cations, among which Mn2+ and Ba2+ were the most effective. When the concentration of both external Ca2+ and Mg2+ was reduced to micromolar amounts, depolarizing photoresponses were observed. In these conditions measurements with radioactive tracers showed a light-modulated efflux of 42K+ or 86Rb+. The light-modulated 42K+ or 86Rb+ efflux was halved by 2 X 6 mM-external K+ and was completely blocked when K+ was raised above 10 mM. These results show that ionic movements through light-sensitive channels are controlled by Ca2+ and Mg2+ and possibly also be the intracellular level of cyclic nucleotides. Moreover, the movement of ions through the light-sensitive channel, does not obey the independence principle.

1-Methyl-3-isobutylxanthine↗

Mechanism of action of the sodium pump in vertebrate photoreceptors.

Membrane potential and photoresponse of rods in the isolated toad retina was recorded while changing the ionic composition of the extracellular medium. In the presence of 10 mM external Cs+ the bright flash response consisted of an initial fast component of about 35 mV followed by a much slower component as large as 50 mV. The slow component of the photoresponse was blocked by micromolar amounts of both ouabain and strophanthidin. The effects induced by the latter were almost completely irreversible. The effects induced by changing the external concentrations of Na+, K+ and Ca2+ were analysed assuming that the amplitude of the slow component of voltage photoresponse reflects the activity of Na+--K+ pump. It is shown that external Na+, K+ and Ca2+ have the same effect on the amplitude of the slow component as on the Na+-K+ pump activity of other tissues.

Animals↗

Antagonism between steady light and phosphodiesterase inhibitors on the kinetics of rod photoresponses.

Treatment of toad rods with phosphodiesterase inhibitors (3-isobutyl-1-methylxanthine, caffeine, theophylline, papaverine, and RO 20-1724) modifies the properties of the intracellular voltage responses to dim flashes of light. 3-Isobutyl-1-methylxanthine at 1-20 microM causes an increase in flash sensitivity and a slowing down of the kinetics of the photoresponses. When the drug concentration is greater than 20 microM, rods also show supralinear behavior, whereby doubling the intensity of a dim flash may increase the response by greater than 2-fold. Sensitivity, kinetics, and supralinear behavior can be restored to normal by steady background illumination while still in the presence of 3-isobutyl-1-methylxanthine. However, the intensity of the steady light needed to restore the sensitivity to control levels is not sufficient to accelerate the kinetics back to control values. The antagonism between the effects of 3-isobutyl-1-methylxanthine and the effects of background illumination is explained by assuming that: (i) the length of time to peak voltage responses to dim flashes of light is inversely proportional to the rate of a chemical reaction; (ii) the rate of this reaction is controlled by an enzyme that is inhibited competitively by 3-isobutyl-1-methylxanthine with a Ki of 3 x 10(-6) M; and (iii) the concentration of a cofactor of this reaction increases proportionally with the intensity of the background illumination.

1-Methyl-3-isobutylxanthine↗

The contribution of the electrogenic sodium-potassium pump to the electrical activity of toad rods.

1. The membrane potential of rods in the isolated toad retina was recorded while changing the ionic composition of the extracellular medium.2. Caesium (Cs(+)) at a concentration of 1 mM was sufficient to completely block the sag from the peak to the plateau in the bright-flash voltage response.3. In the presence of 10 mM-Cs(+) the bright-flash response increased in amplitude to about 90 mV, thus reaching an absolute membrane potential of between -110 and -135 mV. These responses consisted of an initial fast component of about 35 mV followed by a much slower component which could be as large as 50 mV.4. At the peak of the initial fast component the rod membrane conformed closely to the behaviour of a K(+) electrode with a P(Na)/P(K) ratio of 0.023. On average the amplitude of the slow component was about 35 mV in the presence of 2.6 mM-K(+) and was reduced to about 25 mV in a K(+)-free Ringer.5. Addition of 100 muM-strophanthidin to the perfusate induced several reversible changes in the electrical activity of rods. The dark resting membrane potential depolarized by about 5 mV and the kinetics of the voltage response to dim flashes of light slowed down. The voltage sensitivity initially increased by about 30%, but the peak of the response to a bright flash of light was reduced by about 13 mV.6. In rods treated with 10 mM-Cs(+) the slow component present in the bright flash response was abolished by strophanthidin with an apparent K(m) of 3 muM.7. The amplitude of the slow component decreased with a time lag of about 2 min when external Na(+) was reduced. A previous exposure of the retina to a Na(+)-free Ringer solution for at least 3 min modified the voltage photoresponse in a way similar to that observed in the presence of 100 muM-strophanthidin.8. When external Ca(2+) concentration ([Ca(2+)](o)) was increased from 2 to 5 mM the slow component decreased by about 30%. When [Ca(2+)](o) was reduced the slow component increased. A twofold increase was observed when [Ca(2+)](o) was lower than 10(-4) M.9. It is suggested that the slow component of the voltage response in the presence of external Cs(+) is caused by an electrogenic current driven by the Na(+)-K(+) transport system, during a voltage-dependent block of external Cs(+) of some K(+) channels.

Animals↗

Rod photoresponses in the absence of external sodium in retinae treated with phosphodiesterase inhibitors.

Exposure of isolated toad retinae to phosphodiesterase inhibitors, induced changes in the ionic permeability of rod cells. Under similar conditions intracellularly recorded light responses were observed also in the absence of external Na+. Hyperpolarizing photoresponses in Na+-free media required the presence of divalent cations among which Mg2+, Mn2+ and Ba2+ were the most effective.

1-Methyl-3-isobutylxanthine↗

The sodium current underlying the responses of toad rods to light.

1. Intracellular responses were recorded from single rods in the retina of the toads Bufo bufo and Bufo marinus during exposure to solutions in which sodium was replaced by equimolar amounts of choline. 2. Upon moderate reduction (80 and 50 mM) of the external sodium the size of responses to bright flashes decreased as a consequence of both an increase in the resting potential and a decrease of the membrane potential at the peak, while the level of the plateau remained fairly constant. 3. Upon reduction of the external sodium to 22 mM or less, rods hyperpolarized to about the plateau level and failed to respond to illumination. Under these circumstances, membrane depolarization induced by an increased external potassium did not restore the cell responsiveness. Addition of 2-5 mM caesium hyperpolarized the membrane and partially restored the photoresponse. 4. Complete replacements of external sodium with potassium depolarized the rod by 40 +/- 10 mV, and no voltage responses to light could be detected. 5. In the presence of caesium, a nearly complete blockage of the photoresponses was obtained when the external sodium was 5 mM or less. Further reductions of the external sodium did not invert the photoresponses. Application of caesium when the external sodium was nominally zero induced a transient hyperpolarization followed by a slow decay. 6. During exposure to steady illumination, the dependence of the plateau level on the external sodium slowly increased. 7. These results indicate that the ionic current which is directly modulated by the light depends primarily on the external sodium. They suggest also that the current associated with the voltage- and time-dependent process responsible for the sag from peak to plateau of the response to a bright flash of light may have multiple components.

Animals↗

Effects of changing external potassium and chloride concentrations on the photoresponses of Bufo bufo rods.

1. Intracellular responses to light were recorded from Bufo bufo rods in different ionic media. 2. The exposure of the retina to high external [K+] depolarized the rod and modified the time course of the photoresponse. The prominent initial transient of rod responses to bright flashes was drastically reduced in 5 mM-external [K+] and completely disappeared in 26 mM. In high external [K+] the kinetics of responses to dim flashes were considerably slower than in control conditions. 3. When external [Cl-] was changed from 120.6 to 10.6 mM the resting membrane potential decreased and the size of photoresponses increased. Changes in the kinetics similar to those described in high external [K+] were also observed. 4. In many cases exposure of the retina to low external [Cl-] induced oscillations of the resting membrane potential that sometimes became sustained. This instability of the membrane completely disappeared upon restoring to normal conditions. 5. The present results may be explained by assuming the existence of a voltage- and time-dependent conductance active near the dark level of membrane potential. This hypothesis can be represented by an equivalent electrical circuit that includes an inductance (Detwiler, Hodgkin & McNaughton, 1980).

Animals↗

The responses of amacrine cells to light and intracellularly applied currents.

1. Intracellular responses to light were recorded from amacrine cells in the retina of the turtle Pseudemys scripta elegans. 2. The recorded responses were identified on the basis of physiological criteria reported previously (Marchiafava, 1976). Amacrine cells produced transient 'on' and 'off' depolarizing responses irrespective of the retinal area illuminated and of wavelength. 3. The transient depolarizing responses increased by enlarging the illuminated circle up to 120 micrometer in radius. Circles covering larger areas, up to 200 micrometer, produced a relative decrease of the response amplitude. Thus, amacrine cells' receptive fields appear as a central 'excitatory' area of about 120 micrometer radius, surrounded by a 'suppressor' area. 4. Amacrine cells' photoresponses were associated with an increase in membrane conductance. The responses to illumination of central or peripheral areas of the receptive field, however, showed different reversal potentials. The responses to peripheral illumination reversed at about 15 mV above resting potential, while the equilibrium potential of the centre-photoresponses was indicated by extrapolation at about +30 mV. No conductance chance was detectable during steady lights. 5. Repetitive stimulation of the optic nerve invariably reduced amacrine cells' photoresponses, but not those recorded from bipolar cells. It follows then that only ganglion cell photoresponses originating from amacrines' input would be depressed by the nerve stimulation, which thus becomes a reliable test to discriminate whether ganglion cell photoresponses originate from amacrine or bipolar inputs.

Action Potentials↗

Mechanisms of generation of signals in vertebrate photoreceptors.

The electrical responses of rods are analyzed in different ionic environments. It is shown that the dark level of the membrane potential is predominantly determined by a sodium current, while the peak of responses to bright light is controlled by the concentration of external potassium. The sag from the peak to the plateau of photoresponses seems to be generated by different ionic mechanism. The effects produced by substituting the external calcium with EGTA are also analyzed. It is suggested that calcium plays a role in different mechanisms of generation of electrical responses.

Animals↗

Electrical responses of rods in the retina of Bufo marinus.

1. Intracellular responses to flashes and steps of light have been recorded from the outer segment and the cell body of rods in the retina of the Bufo marinus. The identification of the origin of recorded responses has been confirmed by intracellular marking.2. Responses to flashes delivered in darkness or superimposed on a background were analysed. Responses recorded from outer segments conform to the principle of ;spectral univariance'. The shape of the response is not affected by enlarging the spot diameter from 150 to 1000 mum.3. The membrane potential measured in darkness at the outer segments varied from -15 to -25 mV. Injection of steady hyperpolarizing currents increases the size of the response to light; depolarizing currents reduce the response. The mean value of the input resistance is 97 +/- 30 MOmega in darkness and increases by 20-30% during illumination.4. The responses obtained from the cell body of rods have the same shape, time course and spectral sensitivity of those recorded at the outer segment. Injection of steady current at the cell body produces different effects than at the outer segment: hyperpolarizing currents reduce the amplitude of the response to light; depolarizing currents increase the response.5. The experimental data are fitted according to a model similar to that used to describe the responses of turtle cones (Baylor & Hodgkin, 1974; Baylor, Hodgkin & Lamb, 1974a, b).6. The model reproduces the electrical responses of the rod outer segment to a variety of stimuli: (a) brief flashes and steps of light in dark adapted conditions; (b) bright flashes superimposed on background illuminations; (c) pairs of flashes delivered at different time intervals. Responses to hyperpolarizing steps of current are also reproduced by the model.

Animals↗