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Content and Fluxes of Electrolytes

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Abstract

The sections in this article are:

1 null
2 Physicochemical Models of Cellular Behavior
2.1 Prevailing Assumptions
2.2 Osmotic Properties
2.3 Membrane Diffusion
2.4 Electrochemical Potentials
3 Experimental Approaches
3.1 Chemical Dissection
3.2 Isotope Flux Analyses
3.3 Electron Probe Analyses
4 Ionic Transport Mechanisms
4.1 Simple Diffusion or Permeation
4.2 Active Transport
4.3 Exchange Diffusion
5 Control of Cellular H2O
5.1 Osmotic Balance
5.2 Transport Mechanisms
6 Temperature Dependence
6.1 Activation Energy
6.2 Phase Transitions
7 Effects of Excitatory Agents
7.1 Membrane Potentials
7.2 Ionic Fluxes
8 Divalent Ions
8.1 Calcium
8.2 Magnesium
9 Conclusion
Figure 1. Figure 1.

Distribution of extracellular markers in dog carotid arteries. 60Co‐labeled EDTA (•) determined at various times. Horizontal lines represent steady‐state distribution of sucrose and inulin achieved at 3 h incubation. Data are means ± SE from 10 dogs.

From Jones and Swain
Figure 2. Figure 2.

Washout curve of 42K from a rat aorta after in vivo labeling of the various compartments. Logarithm of radioactivity is plotted against time. Graphical analysis individualizes a slow component (half‐renewal time, 70 min) and a fast slope with a half‐time of 2 min.

From Hagemeijer et al.
Figure 3. Figure 3.

Efflux curves of 36Cl at 37°C in normal Krebs, in Krebs‐nitrate, and in normal Krebs + cyanide + iodoacetate. Residual component has been subtracted from the curves. Fast phase obtained after subtraction of slow phase from the curve at 37°C is shown at right. Values are means of 7, 6, and 5 experiments, respectively.

From Villamil et al.
Figure 4. Figure 4.

Analysis of 24Na efflux curve. Points representing radioactivity in the tissue in mmol/kg wet wt (solid circles) can be considered to fall on a straight line after 15 min. Middle component of the curve (open circles) was obtained by subtracting the extrapolated line for the slow component from the experimental values. Values of this second component fell on a straight line after about 5 min, and subtraction of this line from the earlier values gives the third and fastest component (open triangles), the values for which all fell on a straight line. Slopes of the three lines represent the rate constants k1, k2, and k3 for the three components A, B, and C obtained by extrapolation of the lines to zero time. Efflux curve can thus be described by the equation 24Na, mmol/kg wet wt = . Curve shown is a mean curve for 6 muscles. Abscissa, time in min; ordinate, tissue radioactivity in mmol/kg wet wt.

From Wahlström
Figure 5. Figure 5.

representative washout of 24Na from rabbit pulmonary artery, PA (•), and portal vein, PV (▪), in presence of K (5 mM). Logarithm of radioactivity plotted against time. Time scale is expanded at 50 min when the tissue was switched from 1°C to 37°C (arrow).

representative washout of 24Na plotted as in A. Washout was conducted in the presence of ouabain (10−5 M)

From Jones and Miller
Figure 6. Figure 6.

Effects of high KCl solution on the distribution of water in rabbit aorta (8 animals). Abscissa, time in min after transfer to the high KCl solution. Ordinate, water components with total water (•), 60Co‐labeled EDTA space (▴), and smooth muscle (▪) in percent weight. Measurements of tissues remaining in normal Krebs solution are denoted by (○), (▵), and (□) for the respective components. Mean ± SE represented by vertical bars, except when less than the size of the symbol. Points are joined by straight lines.

From Jones et al.
Figure 7. Figure 7.

Ouabain‐dependent (10−4 M) 24Na efflux (•) and 42K influx (▪) into carotid artery from the rabbit at various [K]o. Ratio of the 24Na to 42K flux (▴) appears on top. Arrows indicate the [K]o required for half‐saturation. Curves were computed from Equation . Means ± SE are plotted for 8–12 rabbits.

From Heidlage and Jones
Figure 8. Figure 8.

Changes of membrane potential (mV) and tension development (arbitrary units) of arterial smooth muscle cells as a function of time (min). At time 0 the cells are exposed to a Krebs solution containing 10−5 M ouabain. After 20 min, normal Krebs solution is readmitted for 60 min. Cells are then exposed to K‐free solution for 105 min. Readmission of K (n‐Krebs) is followed after 1 min by addition of ouabain to the solution. After another 1 min the ouabain is washed out again.

From Hendrickx and Casteels
Figure 9. Figure 9.

K‐stimulated 24Na efflux at various [Na]cell for control (•) and Doca‐hypertensive rat aortas (○). [K] of 10 mM was used for stimulation. Arrows indicate [Na]cell required for half‐saturation. Curves are computed from Equation . Because [Na]cell could not be precisely controlled, both variables are means ± SE; n = 5−7 rats. Doca, deoxycorticosterone acetate; [Na]cell, cellular concentration of Na; C, controls.

From Jones
Figure 10. Figure 10.

Effect of alkali metal cations on the rate of loss of 24Na into MgCl2 washout solution. During time indicated by bar, the washout solution contained 46 mM alkali metal cation, replacing some of the Mg. All solutions contained ouabain (2 × 10−5 M) except the K+ ‐ and Rb+ ‐containing solutions (which contained 10−3 M ouabain). Effectiveness in stimulating the 24Na efflux was Na > Li > K > Rb > Cs.

From Brading
Figure 11. Figure 11.

Smooth muscle water at various osmolarities in guinea pig taenia coli. Ordinate: experimental values of cellular H2O in high KCl (•) and normal Krebs solution (▵) are mean ± SE; DS, dry solids. Abscissa: concentration of incubating solution is in mosmol. Seven guinea pigs were used for each type of solution. Dry wt were corrected for the weight of the sucrose in the extracellular space. Solid curve was calculated from Equation , whereas dashed curve was calculated from Equation .

From Jones et al.
Figure 12. Figure 12.

Changes in ion content and tissue weight with time on exposure to K‐free solution (○) or K‐free solution + 5 mM La (▵). Points are means of 4–6 determinations ± SE.

From Brading and Widdicombe
Figure 13. Figure 13.

Arrhenius plot of log rate constants for 42K efflux (•), K‐dependent 24Na efflux (▴), and 24Na efflux into a K‐free solution (▪). Abscissa is the absolute T−1; the equivalent temperature (°C) is portrayed for comparison. Straight lines were fitted by regression techniques. Means ± SE for 4–12 rat aortas are plotted.

Adapted from Kamm et al.
Figure 14. Figure 14.

Typical experiment showing effect of 10−5 M norepinephrine on rate constant of the efflux of 42K (A), 36Cl (B), and 24Na (C) from rabbit ear artery. Loading period in the radioactive solutions for 42K experiments was 3 h, and for 36Cl experiments 1 h. For studying 24Na efflux, K‐depleted cells were loaded for 1 h. Duration of exposure to norepinephrine is represented by bar. Initial high rate constant is due to efflux of radioactivity from extracellular compartment.

From Droogmans et al.
Figure 15. Figure 15.

Effects of norepinephrine (N‐epi) on fraction of aortic 42K exchanged/min from controls (•) and deoxycorticosterone acetate (Doca) hypertensives (○). Points are jointed by straight lines, and a representative standard error (mean ± SE) is indicated by vertical bars. Horizontal bars show period of exposure to norepinephrine at indicated doses.

From Jones et al. , by permission of the American Heart Association
Figure 16. Figure 16.

45Ca desaturation from rabbit aortic strips after 1‐h incubation with 45Ca in presence and absence of extracellular calcium. Values represent mean of 4–8 experiments. A: 45Ca efflux into Ca‐free Ringer's solution. B: effect of adding 1.5 mM calcium on 45Ca efflux during 60–125 min washout interval. C: effect of removal of calcium on 45Ca efflux during 60‐ to 125‐min washout interval.

From Hudgins and Weiss
Figure 17. Figure 17.

Uptake of 45Ca into the La‐resistant Ca fraction of rat aorta. Aortas preincubated for 2 h in physiological solution were transferred to 45Ca physiological solution containing 10−5M norepinephrine with or without preincubation in 45Ca solution without norepinephrine. After various periods of time, aortas were washed for 5 min in 50 mM La solution. Abscissa, time (min) in the radioactive solution. Ordinate, 45Ca content of the La‐resistant Ca fraction (in mmol/kg wet wt) of controls in 45Ca physiological solution (•) of aortas transferred to 45Ca physiological solution containing 10−5 M norepinephrine, without preincubation in absence of norepinephrine (▪) with a preincubation of 5 min (○). Means ± SE shown when they exceeded diameter of the symbol.

From Godfraind
Figure 18. Figure 18.

Effect of sequential norepinephrine (NE) exposures on 45Ca efflux from rabbit aorta. Dots, exposure to NE (10−5 M) for 8 min at 36 min and again for 8 min at 50 min. Squares (control) exposed to NE (10−5 M) for first time at 50 min. Ordinate, rate of 45Ca loss as μmol 45Ca/kg min. Abscissa, time in min. Each point is average of 4 observations.

From Deth and van Breemen


Figure 1.

Distribution of extracellular markers in dog carotid arteries. 60Co‐labeled EDTA (•) determined at various times. Horizontal lines represent steady‐state distribution of sucrose and inulin achieved at 3 h incubation. Data are means ± SE from 10 dogs.

From Jones and Swain


Figure 2.

Washout curve of 42K from a rat aorta after in vivo labeling of the various compartments. Logarithm of radioactivity is plotted against time. Graphical analysis individualizes a slow component (half‐renewal time, 70 min) and a fast slope with a half‐time of 2 min.

From Hagemeijer et al.


Figure 3.

Efflux curves of 36Cl at 37°C in normal Krebs, in Krebs‐nitrate, and in normal Krebs + cyanide + iodoacetate. Residual component has been subtracted from the curves. Fast phase obtained after subtraction of slow phase from the curve at 37°C is shown at right. Values are means of 7, 6, and 5 experiments, respectively.

From Villamil et al.


Figure 4.

Analysis of 24Na efflux curve. Points representing radioactivity in the tissue in mmol/kg wet wt (solid circles) can be considered to fall on a straight line after 15 min. Middle component of the curve (open circles) was obtained by subtracting the extrapolated line for the slow component from the experimental values. Values of this second component fell on a straight line after about 5 min, and subtraction of this line from the earlier values gives the third and fastest component (open triangles), the values for which all fell on a straight line. Slopes of the three lines represent the rate constants k1, k2, and k3 for the three components A, B, and C obtained by extrapolation of the lines to zero time. Efflux curve can thus be described by the equation 24Na, mmol/kg wet wt = . Curve shown is a mean curve for 6 muscles. Abscissa, time in min; ordinate, tissue radioactivity in mmol/kg wet wt.

From Wahlström


Figure 5.

representative washout of 24Na from rabbit pulmonary artery, PA (•), and portal vein, PV (▪), in presence of K (5 mM). Logarithm of radioactivity plotted against time. Time scale is expanded at 50 min when the tissue was switched from 1°C to 37°C (arrow).

representative washout of 24Na plotted as in A. Washout was conducted in the presence of ouabain (10−5 M)

From Jones and Miller


Figure 6.

Effects of high KCl solution on the distribution of water in rabbit aorta (8 animals). Abscissa, time in min after transfer to the high KCl solution. Ordinate, water components with total water (•), 60Co‐labeled EDTA space (▴), and smooth muscle (▪) in percent weight. Measurements of tissues remaining in normal Krebs solution are denoted by (○), (▵), and (□) for the respective components. Mean ± SE represented by vertical bars, except when less than the size of the symbol. Points are joined by straight lines.

From Jones et al.


Figure 7.

Ouabain‐dependent (10−4 M) 24Na efflux (•) and 42K influx (▪) into carotid artery from the rabbit at various [K]o. Ratio of the 24Na to 42K flux (▴) appears on top. Arrows indicate the [K]o required for half‐saturation. Curves were computed from Equation . Means ± SE are plotted for 8–12 rabbits.

From Heidlage and Jones


Figure 8.

Changes of membrane potential (mV) and tension development (arbitrary units) of arterial smooth muscle cells as a function of time (min). At time 0 the cells are exposed to a Krebs solution containing 10−5 M ouabain. After 20 min, normal Krebs solution is readmitted for 60 min. Cells are then exposed to K‐free solution for 105 min. Readmission of K (n‐Krebs) is followed after 1 min by addition of ouabain to the solution. After another 1 min the ouabain is washed out again.

From Hendrickx and Casteels


Figure 9.

K‐stimulated 24Na efflux at various [Na]cell for control (•) and Doca‐hypertensive rat aortas (○). [K] of 10 mM was used for stimulation. Arrows indicate [Na]cell required for half‐saturation. Curves are computed from Equation . Because [Na]cell could not be precisely controlled, both variables are means ± SE; n = 5−7 rats. Doca, deoxycorticosterone acetate; [Na]cell, cellular concentration of Na; C, controls.

From Jones


Figure 10.

Effect of alkali metal cations on the rate of loss of 24Na into MgCl2 washout solution. During time indicated by bar, the washout solution contained 46 mM alkali metal cation, replacing some of the Mg. All solutions contained ouabain (2 × 10−5 M) except the K+ ‐ and Rb+ ‐containing solutions (which contained 10−3 M ouabain). Effectiveness in stimulating the 24Na efflux was Na > Li > K > Rb > Cs.

From Brading


Figure 11.

Smooth muscle water at various osmolarities in guinea pig taenia coli. Ordinate: experimental values of cellular H2O in high KCl (•) and normal Krebs solution (▵) are mean ± SE; DS, dry solids. Abscissa: concentration of incubating solution is in mosmol. Seven guinea pigs were used for each type of solution. Dry wt were corrected for the weight of the sucrose in the extracellular space. Solid curve was calculated from Equation , whereas dashed curve was calculated from Equation .

From Jones et al.


Figure 12.

Changes in ion content and tissue weight with time on exposure to K‐free solution (○) or K‐free solution + 5 mM La (▵). Points are means of 4–6 determinations ± SE.

From Brading and Widdicombe


Figure 13.

Arrhenius plot of log rate constants for 42K efflux (•), K‐dependent 24Na efflux (▴), and 24Na efflux into a K‐free solution (▪). Abscissa is the absolute T−1; the equivalent temperature (°C) is portrayed for comparison. Straight lines were fitted by regression techniques. Means ± SE for 4–12 rat aortas are plotted.

Adapted from Kamm et al.


Figure 14.

Typical experiment showing effect of 10−5 M norepinephrine on rate constant of the efflux of 42K (A), 36Cl (B), and 24Na (C) from rabbit ear artery. Loading period in the radioactive solutions for 42K experiments was 3 h, and for 36Cl experiments 1 h. For studying 24Na efflux, K‐depleted cells were loaded for 1 h. Duration of exposure to norepinephrine is represented by bar. Initial high rate constant is due to efflux of radioactivity from extracellular compartment.

From Droogmans et al.


Figure 15.

Effects of norepinephrine (N‐epi) on fraction of aortic 42K exchanged/min from controls (•) and deoxycorticosterone acetate (Doca) hypertensives (○). Points are jointed by straight lines, and a representative standard error (mean ± SE) is indicated by vertical bars. Horizontal bars show period of exposure to norepinephrine at indicated doses.

From Jones et al. , by permission of the American Heart Association


Figure 16.

45Ca desaturation from rabbit aortic strips after 1‐h incubation with 45Ca in presence and absence of extracellular calcium. Values represent mean of 4–8 experiments. A: 45Ca efflux into Ca‐free Ringer's solution. B: effect of adding 1.5 mM calcium on 45Ca efflux during 60–125 min washout interval. C: effect of removal of calcium on 45Ca efflux during 60‐ to 125‐min washout interval.

From Hudgins and Weiss


Figure 17.

Uptake of 45Ca into the La‐resistant Ca fraction of rat aorta. Aortas preincubated for 2 h in physiological solution were transferred to 45Ca physiological solution containing 10−5M norepinephrine with or without preincubation in 45Ca solution without norepinephrine. After various periods of time, aortas were washed for 5 min in 50 mM La solution. Abscissa, time (min) in the radioactive solution. Ordinate, 45Ca content of the La‐resistant Ca fraction (in mmol/kg wet wt) of controls in 45Ca physiological solution (•) of aortas transferred to 45Ca physiological solution containing 10−5 M norepinephrine, without preincubation in absence of norepinephrine (▪) with a preincubation of 5 min (○). Means ± SE shown when they exceeded diameter of the symbol.

From Godfraind


Figure 18.

Effect of sequential norepinephrine (NE) exposures on 45Ca efflux from rabbit aorta. Dots, exposure to NE (10−5 M) for 8 min at 36 min and again for 8 min at 50 min. Squares (control) exposed to NE (10−5 M) for first time at 50 min. Ordinate, rate of 45Ca loss as μmol 45Ca/kg min. Abscissa, time in min. Each point is average of 4 observations.

From Deth and van Breemen
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Allan W. Jones. Content and Fluxes of Electrolytes. Compr Physiol 2011, Supplement 7: Handbook of Physiology, The Cardiovascular System, Vascular Smooth Muscle: 253-299. First published in print 1980. doi: 10.1002/cphy.cp020211