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PubMed1975

Estimation of surface charges in some biological membranes.

The resting membrane potential data existing in the literature for the giant axon of the squid, frog muscle and barnacle muscle have been analyzed from the standpoint of the theory of membrane potential due to Kobatake and co-workers. The average values derived for the effective charge density phi chi (where phi is a constant, 0 less than phi less than 1, and represents the fraction of counterions that are free, and chi is the stoichiometric charge density in the membrane) present on the different biomembranes existing in their normal ionic environment are 0.3, 0.325 and 0.17 M for the squid axon, frog and barnacle muscles, respectively. On the assumption that the values of phi are 0.4 and 0.2 for nerve and muscle membranes, respectively, values of 0.75, 1.62 and 0.85 M have been derived for the stoichiometric charge density (chi) present in the respective biological membranes. These correspond to 1 negative charge per 222, 103 and 195 A2 of the membrane area of the squid axon, frog and barnacle muscles, respectively.

Animals↗
PubMed1985

A minimum mechanism for Na+-Ca++ exchange: net and unidirectional Ca++ fluxes as functions of ion composition and membrane potential.

Both simultaneous and consecutive mechanisms for Na+-Ca++ exchange are formulated and the associated systems of steady-state equations are solved numerically, and the net and unidirectional Ca++ fluxes computed for a variety of ionic and electrical boundary conditions. A simultaneous mechanism is shown to be consistent with a broad range of experimental data from the squid giant axon, cardiac muscle and isolated sarcolemmal vesicles. In this mechanism, random binding of three Na+ ions and one Ca++ on apposing sides of a membrane are required before a conformational change can occur, translocating the binding sites to the opposite sides of the membranes. A similar (return) translocation step is also permitted if all the sites are empty. None of the other states of binding can undergo such translocating conformational changes. The resulting reaction scheme has 22 reaction steps involving 16 ion-binding intermediates. The voltage dependence of the equilibrium constant for the overall reaction, required by the 3:1 Na+: Ca++ stoichiometry was obtained by multiplying and dividing, respectively, the forward and reverse rate constants of one of the translocational steps by exp(-FV/2RT). With reasonable values for the membrane density of the enzyme (approximately 120 sites micron 2) and an upper limit for the rate constants of both translocational steps of 10(5) . sec-1, satisfactory behavior was obtainable with identical binding constants for Ca++ on the two sides of the membrane (10(6) M-1), similar symmetry also being assumed for the Na+ binding constant (12 to 60 M-1). Introduction of order into the ion-binding process eliminates behavior that is consistent with experimental findings.

Animals↗
PubMed1983

Effects of an outward water flow on potassium currents in a squid giant axon.

The excitation of the squid giant axon was analyzed under an outward water flow through the membrane product by an osmotic gradient. The outward water flow made an undershoot of the action potential larger by about 25 mV without decreasing its peak largely. It also made EK more negative but not ENa. The effect of the outward water flow was specific for the potassium channel. The outward current increased and its decline during a long-lasting depolarization became less prominent under the outward water flow. At the same time, inward tail currents for the potassium channel decreased extraordinarily without a large change in the time course. The potassium conductance had a marked rectification in the direction of the water flow. The undershoot of the action potential under the outward water flow was very sensitive to potassium ions in the external solution. Eight mM KCl was effective to diminish the undershoot and to restore the change in EK by about 60% but gave no effect on the reduced tail current. The outward water flow effect can be explained not only by the change in a local concentration of potassium ions at the mouths of the potassium channel due to a sieving but also by the rectification in a hydrodynamic manner.

Action Potentials↗
PubMed1981

Improvements in optical methods for measuring rapid changes in membrane potential.

In an effort to increase the utility of optical methods for measuring membrane potential in excitable cells, an additional 369 dyes were tested on giant axons from the squid. Several promising dyes with relatively large absorption and fluorescence signals are described. In addition, a simple modification of the apparatus led to a sixfold increase in the size of dye-related birefringence signals. In preparations with a suitable geometry, these signals are as large as absorption signals but photodynamic damage and bleaching are eliminated when wavelengths longer than the absorption band are used.

Animals↗
PubMed1981

Lipid vesicle-mediated alterations of membrane cholesterol levels: effects on Na+ and K+ currents in squid axon.

We demonstrate that cholesterol can exchange from sonicated lipid vesicles to a perfused squid axon membrane and that vesicles with varying cholesterol/phospholipid (C/P) mole ratios can be used to achieve either net loading or net depletion of axon membrane cholesterol. Two types of evidence were obtained which show that net loading or depletion of cholesterol was achieved: (i) changes in the cholesterol/phospholipid (C/P) mole ratios of axons, and (ii) visualization of cholesterol depleted from the preparation by cholesterol-free vesicles by thin-layer chromatography. The C/P mole ratios indicate that cholesterol levels in the preparation were increased or decreased by 30-40%. Increasing or decreasing membrane cholesterol levels were ineffective in altering the Na+ or K+ occurrents in voltage-clamped axons. In addition, we determined that cholesterol "flip-flop" across the axonal membrane occurred with a t 1/2 of 7.3 to 15.3 min.

Animals↗
PubMed1984

Effects of D-600 on sodium current in squid axons.

The effects of the calcium antagonist D-600 (methoxyverapamil) on the excitatory inward sodium current, INa, of internally perfused squid giant axons were studied under voltage-clamp conditions. We observed little or no effect of the drug when it was added to the external solution at concentrations of 10-200 microM. Furthermore, it did not produce a frequency, or use-dependent block of INa when repetitive voltage-clamp pulses were used at rates of 2-5 Hz. However, it did produce use-dependent blockade of INa when it was placed internally at a concentration of 200 microM. These results in conjunction with other studies suggest that D-600 is a selective blocker of calcium channels in squid axons when the drug is placed in the external solution. Its effects, when placed in the internal solution, are similar to those of permanently charged local anesthetic derivatives, which also produce use-dependent block of INa.

Animals↗
PubMed1988

Effects of arachidonic acid and the other long-chain fatty acids on the membrane currents in the squid giant axon.

The effects of arachidonic acid and some other long-chain fatty acids on the ionic currents of the voltage-clamped squid giant axon were investigated using intracellular application of the test substances. The effects of these acids, which are usually insoluble in solution, were examined by using alpha-cyclodextrin as a solvent, alpha-cyclodextrin itself had no effect on the excitable membrane. Arachidonic acid mainly suppresses the Na current but has little effect on the K current. These effects are completely reversed after washing with control solution. The concentration required to suppress the peak inward current by 50% (ED50) was 0.18 mM, which was 10 times larger than that of medium-chain fatty acids like 2-decenoic acid. The Hill number was 1.5 for arachidonic acid, which is almost the same value as for medium-chain fatty acids. This means that the mechanisms of the inhibition are similar in both long- and medium-chain fatty acids. When the long-chain fatty acids were compared, the efficacy of suppression of Na current was about the same value for arachidonic acid, docosatetraenoic acid and docosahexaenoic acid. The suppression effects of linoleic acid and linolenic acid on Na currents were one-third of that of arachidonic acid. Oleic acid had a small suppression effect and stearic acid had almost no effect on the Na current. The currents were fitted to equations similar to those proposed by Hodgkin and Huxley (Hodgkin, A.L., Huxley, A.F. (1952) J. Physiol (London) 117:500-544) and the change in the parameters of these equations in the presence of fatty acids were calculated.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗
PubMed1978

Action of extracellular pH on Na+ and K+ membrane currents in the giant axon of Loligo vulgaris.

Voltage-clamp currents and resting membrane potential of squid giant axons have been studied at extracellular pH varying between 4 and 10. The membrane currents, analyzed according to the Hodgkin-Huxley equations, showed that sodium permeability, PNa(E), and potassium conductance gK(E), curves were shifted toward positive voltages by different amounts and slightly depressed as the external pH was lowered. Under the same conditions, taum(E) and taun(E) were found to be enhanced and shifted to a larger extent in the same direction. The rate constants alpham and alphan were shifted substantially toward positive voltages, but betam and betan changed hardly at all. The shift of the alpham(E) curve was analyzed in terms of a fixed surface charge model; it indicates that unspecific negative groups with an approximate pKa of 4.5 are located in the vicinity of sodium active sites with an average charge separation of 8 A. A similar figure is obtained for the potassium system from the shift of the alphan(E) curve.

Animals↗
PubMed1978

Effect of calcium upon sodium inactivation in the giant axon of Loligo pealei.

Giant axons of Loligo pealei were voltage clamped in artificial seawater solutions containing varying concentrations of calcium from 10 to 100 mM, and the sodium conductance inactivation was measured with a series of two-pulse experiments. The h infinity vs. voltage curve showed a shift of about 10 mV in the depolarizing direction on the voltage axis for a tenfold increase in external calcium without substantial alteration in the slope of the voltage dependence. The kinetics of the inactivation process were found to be exponential for hyperpolarizing prepulses, but showed some indication of a sigmoidal decay for depolarizing prepulses in all calcium concentrations employed. Increasing calcium increased the delay in the sigmoidal response. The inactivation time constant tauh increased as a function of calcium concentration over the potential range studied, -10 to -90 mV. The values of the rate constants alphah and betah are decreased with an increase in calcium and these effects are not consistent with parallel shifts of the rate constant vs. voltage curves along the voltage axis for changes in calcium concentration. Magnesium does not behave as an equimolar substitute for calcium. The effect of a solution containing 10 mM calcium and 50 mM magnesium is intermediate to that of solutions containing 10 and 30 mM calcium alone. Predictions of a recent model for the sodium conductance (Moore, J.W., Cox, E.B., 1976 Biophys. J. 16:171) which employs calcium binding were compared with the experimental data.

Animals↗
PubMed1985

Are axoplasmic microtubules necessary for membrane excitation?

The excitability of the squid giant axon was studied as a function of transmembrane hydrostatic pressure differences, the latter being altered by the technique of intracellular perfusion. When a KF solution was used as the internal medium, a pressure difference of about 15 cm water had very little effect on either the membrane potential or excitability. However, within a few minutes after introducing either a KCl-containing, a KBr-containing, or a colchicine-containing solution as the internal medium, with the same pressure difference across the membrane, the axon excitability was suppressed. In these cases, removal of the pressure difference restored the excitability, indicating that the structure of membrane was not irreversibly damaged. Electron-microscopic observations of these axons revealed that the perfusion with a KF solution or colchicine-containing solution preserves the submembranous cytoskeletal layer, whereas perfusion with a KCl or KBr solution dissolves it. These results suggest that the submembranous cytoskeletons including microtubules provide an important mechanical support to the excitable membrane but are not essential elements in channel activities.

Action Potentials↗
PubMed1982

Properties of sodium and potassium channels of the squid giant axon far below 0 degrees C.

Squid giant axon could be excited in concentrated glycerol solutions containing normal concentrations of electrolytes, when osmolalities of solutions inside and outside the axon were matched. These glycerol solutions did not freeze at the temperature as low as -19 degrees C. The nerve excitation in these solutions were observed at this low temperature. The excitation process at this low temperature was slowed down and time constants of the excitation kinetics were several hundredfold larger than those in normal seawater at 10 degrees C, under which temperature the squid habituated. The temperature coefficients for the electrophysiological membrane parameters under this condition were larger than those in normal seawater above 0 degrees C. The Q10 value for the conduction velocity was 2.0 and that of the duration of the action potential was around 8.5. The time course of the membrane currents was also slowed with the Q10 value of around 5 and the magnitude decreased with the Q10 value of around 2 as the temperature was lowered. The Q10 values for the kinetics of the on process of the Na-channel were around 4.5 and were almost the same as those of the off process of the Na-channel in the wide range of the temperature below 0 degrees C. The Q10 value of the on process of K-channel was around 6.5 and was larger than those for Na-channel. The Q10 values increased gradually as the temperature was lowered.

Action Potentials↗
PubMed1990

Chloramine-T alters the nerve membrane birefringence response.

The change in birefringence during depolarizing voltage-clamp pulses of internally perfused squid giant axons are biphasic. There is a rapid decrease in birefringence with a 220-microsec half time at 8 degrees C followed by a slow decrease over the next several milliseconds. After the pulse there is a rapid recovery which is smaller than the initial rapid decrease followed by a slow recovery phase. The rate of change of the slow phase during the pulse is more rapid for larger depolarizations. After the pulse the rate of change is more rapid for more negative potentials. 3.6 mM chloramine-T, applied externally until the sodium currents were prolonged and inactivation was removed, removed the slow phase of the birefringence response both during and after the pulse and made the fast 'off' response as large as the fast 'on' response. Two anesthetics reduced the birefringence response by about 20%. A rocking helix model is presented which relates the birefringence findings and earlier gating current experiments.

Animals↗