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L Cleemann

Publications and source records attributed to L Cleemann.

33 records · Page 2Linked to original sources

An ammonia-sensing air gap microelectrode.

An ammonia-sensing air gap microelectrode has been designed on the basis of a neutral carrier pH-sensing inner electrode. This electrode has a tip diameter of 2 to 5 microns, has a simple design, is easy to fabricate, and has a long shelf life. Its response to ammonium is linear in the range 3 x 10(-5) to 10(-2) M and its response time (95%) is 10 to 15 s. The electrode was converted to a microsensor for urea by immobilization of urease within its tip. The linear response to urea ranged from 3 x 10(-4) to 10(-2) M and the response time was 15 to 20 s.

Ammonia↗

Epinephrine enhances Ca2+ current-regulated Ca2+ release and Ca2+ reuptake in rat ventricular myocytes.

The voltage dependence of the intracellular Ca2+ transients was measured in single rat ventricular myocytes with the fluorescent Ca2+ indicator dye fura-2. The whole-cell voltage clamp technique was used to measure the membrane current, and 0.9 mM fura-2 was loaded into the cell by including it in the dialyzing solution of the patch electrode. A mechanical light chopper operating at 1200 Hz was used to obtain simultaneous measurements of the intracellular Ca2+ activity with fluorescence excitation on either side of the isosbestic point (330 nm and 410 nm). The symmetry of the two optical Ca2+ signals was used as a criterion to guard against artifacts resulting, for instance, from motion. The voltage dependence of peak Ca2+ current and the Ca2+ transient measured 25 ms after depolarizing clamps from a holding potential of -40 mV were bell-shaped and virtually identical. The Ca2+ entry estimated from the integral of the Ca2+ current (0 mV, 25 ms) corresponds to a 5-10 microM increase in the total intracellular Ca2+ concentration, whereas the optical signal indicated a 100 microM increase in total intracellular Ca2+. Repolarization of clamp pulses from highly positive potentials were accompanied by a second Ca2+ transient, the magnitude of which, when summed with that measured during depolarization, was nearly constant. Ryanodine (10 microM) had little or no effect on the peak Ca2+ current but reduced the magnitude of the early Ca2+ transients by 70-90%. Epinephrine (1 microM) increased the Ca2+ current and the Ca2+ transients, accelerated the rate of decline of the Ca2+ transients at potentials between -30 and +70 mV, and reduced the intracellular [Ca2+] below baseline at potentials positive to +80 or negative to -40 mV, where clamp pulses did not elicit any Ca2+ release. Elevation of intracellular cAMP mimicked the relaxant effect of epinephrine at depolarizing potentials, whereas elevation of extracellular [Ca2+] did not. These results suggest that most of the activator Ca2+ in rat ventricular cells is released from the sarcoplasmic reticulum as a graded response to sarcolemmal Ca2+ influx. Consistent with a graded Ca2+-induced Ca2+ release we find that epinephrine increases the internal Ca2+ release by increasing the Ca2+ current. Epinephrine may also increase the Ca2+ content of the sarcoplasmic reticulum that may, in turn, increase the Ca2+-induced Ca2+ release. The relaxant effect of epinephrine appears to be caused by enhanced rate of Ca2+ resequestration and is mediated by adenylate cyclase system.

Algorithms↗

Role of Ca2+ channel in development of tension in heart muscle.

Role of Ca2+ Channel in Development of Tension in Heart Muscle. Journal of Molecular and Cellular Cardiology (1987) 19, 527-553. In mammalian and amphibian hearts Ca2+ enters the myocardial cells via voltage-gated Ca2+ channels. The role of Ca2+ channels in transporting the activator Ca2+ was probed by examining the redevelopment of tension and ICa in voltage-clamped myocardial strips after step recovery of Ca2+ channel from photolabile Ca2+ antagonist. Comparison of the kinetics of redevelopment of tension in the frog and mammalian heart following the photoinactivation of Ca2+ channel blockers showed that in the frog heart tension redevelops fully in one beat, while 5 to 7 beats were required for full redevelopment of tension in the mammalian heart. Ca2+ depletion studies using Antipyrylazo III showed that extracellular Ca2+ depletion during the action potential occurs primarily via the Ca2+ channel. In the frog heart sufficient Ca2+ is transported from the extracellular space during a single beat to activate tension. In the mammalian heart ICa triggers the release of internal stores, which when only fully loaded caused full development of tension. ICa also was involved in loading the intracellular Ca2+ pools. Two types of Ca2+ channels have been identified in the mammalian myocytes. The low-threshold rapidly inactivating Ca2+ channels appear to be involved in trigger release of Ca2+, while the high threshold (conventional Ca2+ channels) seem to be involved primarily in reloading of internal Ca2+ pools.

Animals↗

Luminal and basolateral surface membranes of secretory acinar cells: electrophysiological comparison of cationic sensitivities.

Cation sensitivities (K+, Na+, and Ca2+) of luminal and basolateral membrane surfaces of secretory acinar cells were compared using a luminally perfused and externally superfused salivary gland from the aquatic snail, Helisoma trivolvis. Tight junctions delimiting the two membrane surfaces were observed near the acinar lumen suggesting that the total membrane area exposed to the superfusion solution exceeded that in contact with the luminal perfusion solution. The resting membrane potential of acinar cells was found to be dependent upon the K+ concentration in both the external superfusion and the luminal perfusion solutions. Unilateral K+ elevation at either membrane surface produced a rapid and sustained depolarization of the acinar cell. For a given K+ concentration, the level of depolarization produced by K+ elevation at the basolateral surface was significantly higher than at the luminal surface. The highest level of membrane depolarization was observed following simultaneous K+ elevation at both membrane surfaces. The ability of acinar cells to generate overshooting action potentials in response to electrical field stimulation was dependent upon both Na+ and Ca2+. Complete blockade invariably occurred following bilateral removal of either cation. The effects of unilateral removal of either Na+ or Ca2+ proved to be somewhat variable. In general, unilateral removal of Na+ was more effective in reducing the regenerative response than Ca2+ while removal of either cation from the basolateral surface was more effective in reducing the regenerative response than its removal from the luminal surface. Electrically evoked action potentials in acinar cells could also be blocked with unilateral application of the Ca2+ antagonist, cadmium (Cd2+), at either membrane surface. However, higher Cd2+ concentrations were required to achieve complete blockade when applied to the luminal than to the basolateral gland surface. This result fails to support a hypothesis of voltage-sensitive Ca2+ channels being spatially restricted to the luminal cell surface in this preparation.

Action Potentials↗

Optical measurement of voltage-dependent Ca2+ influx in frog heart.

Sarcolemmal Ca2+ movements in frog ventricular strips were measured by monitoring Ca2+ depletion from the extracellular space with an impermeant Ca indicator dye, antipyrylazo III. Ca2+ depletion was measured as a weighted average of light signals recorded simultaneously at three different wavelengths. This weighting procedure was designed to reduce the motion-induced light scattering and to enhance the Ca2+-related optical signals. Comparison of the time course of Ca2+ depletion signal with that of contraction showed that the rate of Ca2+ depletion was maximal immediately after the upstroke of the action potential but prior to the onset of tension. Peak Ca2+ depletion was reached toward the end of the action potential and amounted to a 10-50 microM decrease in the total extracellular Ca2+ concentration. The reaccumulation of extracellular Ca2+ seen after the action potential was 2-5 sec slower than the relaxation of tension. The rate of Ca2+ depletion had a bell-shaped voltage dependence and was enhanced by epinephrine, suggesting that Ca2+ influx occurred primarily through a slowly inactivating ionic channel. Ca2+ transport through the Na+-Ca2+ exchange system was not significantly altered in the presence of strophanthidin or with decrease of extracellular K+ concentration despite marked potentiation of tension by these agents. Ca2+ depletion measured at the end of a 1-sec clamp pulse had a voltage dependence noticeably different from that of the developed tension. This finding may suggest that a fraction of activator Ca2+ is released from membrane-bound Ca2+ pools in a voltage-dependent manner. Our results show that Ca2+ indicator dyes can be used not only to measure rapid changes in the extracellular Ca2+ concentration during contraction, but also to quantify the contribution of various sarcolemmal Ca2+ transport systems to the generation of tension in cardiac muscle.

Action Potentials↗

Optical measurements of extracellular calcium depletion during a single heartbeat.

The impermeant dye antipyrylazo III was used to measure depletion of extracellular calcium and net influx of calcium through the sarcolemma during the cardiac action potential. It was found that calcium entry occurs continuously during the action potential and is under direct control of the membrane potential. The inotropic action of epinephrine is accompanied by increased influx of calcium, while strophanthidin enhances the twitch without altering calcium influx during the action potential.

Action Potentials↗

Measurement of intracellular 42K diffusion in frog ventricular strips.

The longitudinal distribution of 42K was measured in strips of frog ventricular muscle placed in a partitioned perfusion chamber. A radiation detector placed directly under the chamber was moved from point to point to scan the longitudinal distribution of 42K. The detector was focussed on a 0.6 mm segment of the strip by means of two slits and two Geiger tubes. A beta-particle from the strip was counted only if it passed through both Geiger tubes. This arrangement improved the spatial resolution and decreased the background and the sensitivity to Compton electrons. The intracellular diffusion constant measured with this system is 3.7 X 10(-6) cm2/s.

Animals↗

Reduction of the sucrose-saline interdiffusion in the sucrose gap technique by controlled compression of the extracellular space in myocardial preparations.

The time course and the extent of the sucrose-saline interdiffusion in the sucrose gap technique was investigated experimentally. 14C-sucrose diffusion and voltage clamp experiments were performed with designs of the sucrose apparatus allowing accurate control of the size of the holes in the rubber partitions defining the sucrose gap. Experiments with frog ventricular strips demonstrated that compression of the extracellular space to around 15% of its undisturbed value was possible without disturbance of the intracellular conduction pathway. The procedure reduced the sucrose-saline diffusion significantly and proved indispensible for obtaining reliable voltage-current relations in voltage clamp experiments.

Action Potentials↗

Heart muscle. Intracellular potassium and inward-going rectification.

The cellular K content of frog ventricular strips is monitored using 42K. Cellular K loss evoked by cardiac glycosides or a low extracellular K concentration is accompanied by a more than proportional decrease in the conductance of the resting membrane and the rate of rapid repolarization of the action potential. Voltage clamp experiments relate these changes to a decrease in the magnitude of an inward-rectifying K current. Current-voltage relations measured before and after changing the extracellular K concentration cross each other. This violation of the independence principle has previously suggested that extracellular K ions are required to open the rectifier channel (Cleemann and Morad. 1979. J. Physiol. 286: 113). Decreasing the cellular K content decreases the outward membrane current at all membrane potentials by an amount that is consistent with the independence principle. This suggests that the gating process is not sensitive to intracellular K ions. These findings are consistent with a previously published model for inward-going rectification.

Animals↗

Tunicate heart as a possible model for the vertebrate heart.

A number of problems in cardiac physiology are related to the structural complexity of the tissue. For instance, the tortuous and partially confined extracellular space represents a slowly exchangable compartment in which ions or metabolites may accumulate or deplete during activity. Other problems emanate from the inhomogeneous distribution of intracellular potential and the branching nature of the myocardial cells. Our results suggest that the sea potato heart may be functionally treated as a single layer of myofilaments adjacent to a single active membrane which controls excitability and E-C coupling processes. Yet in many ways this simple structure functions similarly to the vertebrate heart when comparing the action potential, the calcium requirement, and the mechanics of muscular contraction. It is concluded, therefore, that the sea potato heart serves as a good model of the vertebrate heart for investigation of the fundamental properties of cardiac muscle. In fact, this preparation may make it possible to examine the details of the molecular mechanisms of ionic transport and mechanics of muscle contraction, thereby supplementing the results that have been or are being obtained from the mammalian myocardium.

Action Potentials↗

Extracellular potassium accumulation in voltage-clamped frog ventricular muscle.

1. Application of voltage clamp pulses (1--10 sec) to frog ventricular strips causes temporary changes in the extracellular K concentration. 2. The changes in the extracellular K concentration can be estimated from (a) slowly decaying post-clamp after-potentials, (b) changes in the action potential duration, and (c) measurements with a K-selective micro-electrode. 3. The depolarization of the resting potential and the shortening of the action potential are present in approximately the same proportions during voltage-clamp induced extracellular K accumulation and during perfusion with a K-ricn Ringer solution but small consistent differences are noticed. 4. The measurements of the after-potential, the action potential shortening, and the K-electrode response were analysed as indicators of extracellular K+ activity and it was concluded that the after-potential provides the most convenient and reliable estimate of the absolute magnitude of the voltage-clamp induced extracellular K accumulation. 5. The depolarizing after-potentials decay more slowly than the hyperpolarizing after-potentials but it is found that this reflects the selectivity of the membrane to K+ concentrations as predicted by the Nernst or the Goldman equations. 6. Analysis of the redistribution of accumulated K+ from the decay of the after-potential suggests that the major part of the redistribution process can be described by a single time constant (2--4 sec). A much longer time constant is required for a smaller component of the 'tail' in order to bring [K]o to the normal resting state. 7. N-shaped relations similar to the 'steady state' current-voltage relation are obtained when the post-clamp after-potential, the action potential shortening, and the K-electrode response are plotted versus the clamped membrane potential. The maxima of these curves are located around -40 mV and the minima around -20 mV. 8. In spite of a significant outward membrane current (1--1.5 microamperemeter) in the minimum region (-20 mV), the post-clamp after-potential is often hyperpolarizing in nature suggesting extracellular K depletion. 9. These findings indicate that the K efflux is lower at -20 mV than at both higher and lower potentials and suggest that the N-shape 'steady state' current-voltage relation mainly reflects the voltage dependency of the K current. 10. A theory for K accumulation in a single compartment is presented which predicts that a simple linear RC-circuit may describe the electrical response of the preparation in a limited potential range around the resting potential. The extracellular accumulation space was estimated to be 13--16% of the total volume of the preparation. It is tentatively suggested that the accumulation space is equivalent to the subendothelial fraction of the extracellular space.

Action Potentials↗

Potassium currents in frog ventricular muscle: evidence from voltage clamp currents and extracellular K accumulation.

1. The single sucrose voltage clamp technique was used to control the membrane potential of strips of frog ventricular muscle and to measure the membrane current. The extracellular K accumulation was estimated from the after-potential observed after the release of the voltage clamp. 2. Comparing the time course of the membrane current to the time course of the development of the after-potential at different membrane potentials, it was found that all slow current changes are related to changes in the K current across the membrane. 3. Based on measurements of membrane current and the after-potential, the total membrane current was separated into two fractions: (a) the K current which gives rise to K accumulation and (b) the residual membrane current which is unrelated to K accumulation. The current-voltage relation for the residual membrane current is linear or slightly inwardly-rectifying. Residual current is zero at the resting potential and increases to about 1 microamperemeter/cm2 at -20 mV. 4. The measured membrane currents and after-potentials indicate qualitative differences between the K currents which dominate below and above -20 mV. More negative to -20 mV the after-potential develops rapidly while at potentials positive to -20 mV the after-potential develops with some delay. 5. The current dominating below -20 mV is inwardly-rectifying. The current-voltage relation has a maximum (about 2 microamperemeter/cm2) and a region with marked negative slope conductance. The outward current in the region of negative slope conductance is increased with increasing [K]o. 6. A model for the inwardly rectifying K current is described. The model accurately reproduces the shape of the measured current-voltage relations and their modification by alterations in the extracellular K concentration. The model is also compatible with the observation that all slow current changes below -20 mV are directly related to K accumulation. 7. The K current which dominates at potentials positive to -20 mV is activated by a potential and time dependent process which is unrelated to extracellular K accumulation. 8. Q10 for the magnitude of the inwardly rectifying K current is about 1.35 while the Q10 for the rate of increase of the time dependent K current is about 3--4. 9. Cs blocks the inwardly recitfying K current but has little effect on the time dependent K current. 10. The changes in the action potential duration caused by increasing the extracellular K concentration or addition of Cs to the perfusate can be explained by the effect of K and Cs on the inwardly rectifying K current.

Animals↗

The inotropic action of adrenaline on cardiac muscle: does it relax or potentiate tension?

Adrenaline has been shown to increase twitch tension and enhance relaxation in cardiac muscle. In mammalian myocardium, a unitary mechanism, namely facilitated uptake of calcium by the sarcoplasmic reticulum, is proposed to increase the internal recirculating store of calcium (thereby potentiating twitch tension) and simultaneously enhance relaxation. In frog ventricular myocardium, where tension is directly controlled by membrane potential, adrenaline seems to produce its positive inotropic effect by increasing the duration and amplitude of cardiac action potential plateau. If adrenaline is prevented from changing the action potential, either by electrical or pharmacologic means, the relaxant effect of the drug is unmasked. The results suggest that in frog ventricle, unlike mammalian myocardium, adrenaline may not have a 'true' positive inotropic effect independent of membrane potential. The findings in frog ventricle, where there is little or no internal recirculation of calcium, are consistent with the model proposed for the mammalian myocardium.

Action Potentials↗

Extracellular potassium accumulation and inward-going potassium rectification in voltage clamped ventricular muscle.

Measurements of afterpotential, action potential duration, and output of a potassium-sensitive microelectrode indicate that the application of long clamp pulses (1 to 8 seconds) to frog ventricular muscle is accompanied by a change in the extracellular potassium concentration. The plot of the magnitude of the potassium accumulation against the clamped membrane potential yields an N-shaped relation similar to the "steady state" current-voltage relation. The accumulation studies confirm a strong inward-going (anomalous) potassium rectification.

Action Potentials↗