Effects of ouabain on background and voltage-dependent currents in canine colonic myocytes

1990 ◽  
Vol 259 (3) ◽  
pp. C402-C408 ◽  
Author(s):  
E. P. Burke ◽  
K. M. Sanders
Keyword(s):  
Membrane Potential ◽  
Potential Gradient ◽  
Circular Muscle ◽  
Input Resistance ◽  
Pump Current ◽  
Muscle Layer ◽  
Membrane Properties ◽  
Resting Potential ◽  
Voltage Dependent ◽  
In Cells

Previous studies have suggested that the membrane potential gradient across the circular muscle layer of the canine proximal colon is due to a gradient in the contribution of the Na(+)-K(+)-ATPase. Cells at the submucosal border generate approximately 35 mV of pump potential, whereas at the myenteric border the pump contributes very little to resting potential. Results from experiments in intact muscles in which the pump is blocked are somewhat difficult to interpret because of possible effects of pump inhibitors on membrane conductances. Therefore, we studied isolated colonic myocytes to test the effects of ouabain on passive membrane properties and voltage-dependent currents. Ouabain (10(-5) M) depolarized cells and decreased input resistance from 0.487 +/- 0.060 to 0.292 +/- 0.040 G omega. The decrease in resistance was attributed to an increase in K+ conductance. Studies were also performed to measure the ouabain-dependent current. At 37 degrees C, in cells dialyzed with 19 mM intracellular Na+ concentration [( Na+]i), ouabain caused an inward current averaging 71.06 +/- 7.49 pA, which was attributed to blockade of pump current. At 24 degrees C or in cells dialyzed with low [Na+]i (11 mM), ouabain caused little change in holding current. With the input resistance of colonic cells, pump current appears capable of generating at least 35 mV. Thus an electrogenic Na+ pump could contribute significantly to membrane potential.

Role of sodium pump in membrane potential gradient of canine proximal colon

1988 ◽  
Vol 254 (4) ◽  
pp. C475-C483 ◽  
Author(s):  
E. P. Burke ◽  
J. B. Reed ◽  
K. M. Sanders
Keyword(s):  
Membrane Potential ◽  
Potential Gradient ◽  
Circular Muscle ◽  
Bath Temperature ◽  
Muscle Layer ◽  
Membrane Potentials ◽  
Resting Potential ◽  
Equilibrium Potential ◽  
Circular Layer ◽  
External K

A large gradient in membrane potential exists through the thickness of the circular layer in canine colonic muscles. This study tested the effects of several experimental manipulations known to block electrogenic sodium pumping on the resting potentials of colonic muscles. Membrane potentials were recorded with microelectrodes from cells through the circular muscle layer. In cells adjacent to the submucosal surface of the circular layer, application of ouabain (10(-6) to 10(-5) M) caused an average membrane depolarization of 36 mV. Removal of the external K+ resulted in depolarizations similar to the effect of ouabain. Readmission of K+ (5.9 mM) produced repolarization and an additional hyperpolarization that averaged 13 mV beyond the resting potential. When exposed to 15 mM K+, cells hyperpolarized well beyond the estimated potassium equilibrium potential (EK). Ouabain blocked the repolarization in response to reintroduction of external K+. Lowering the bath temperature to 20 degrees C rapidly depolarized membrane potential; rewarming repolarized cells. Ouabain and K+-free solutions blocked the repolarization response to rewarming. Cells also depolarized when exposed to solutions in which the NaCl was replaced with LiCl. Membrane potentials of cells within the bulk of the circular layer decreased as a function of distance from the submucosal border. Cells at the myenteric border of the circular muscle were not significantly affected by ouabain and K+-free solution, but these treatments abolished the gradient in membrane potential across the circular layer.(ABSTRACT TRUNCATED AT 250 WORDS)

Perforated patch-clamp analysis of the passive membrane properties of three classes of hippocampal neurons

1992 ◽  
Vol 67 (3) ◽  
pp. 508-529 ◽  
Author(s):  
N. Spruston ◽  
D. Johnston
Keyword(s):  
Membrane Potential ◽  
Hippocampal Neurons ◽  
Pyramidal Neurons ◽  
Physiological Saline ◽  
Membrane Properties ◽  
Resting Potential ◽  
Voltage Dependent ◽  
Perforated Patch Clamp ◽  
Passive Membrane Properties ◽  
Passive Membrane

1. Perforated patch-clamp recordings were made from the three major classes of hippocampal neurons in conventional in vitro slices prepared from adult guinea pigs. This technique provided experimental estimates of passive membrane properties (input resistance, RN, and membrane time constant, tau m) determined in the absence of the leak conductance associated with microelectrode impalement or the washout of cytoplasmic constituents associated with conventional whole-cell recordings. 2. To facilitate comparison of our data with previous results and to determine the passive membrane properties under conditions as physiological as possible, recordings were made at the resting potential, in physiological saline, and without any added blockers of voltage-dependent conductances. 3. Membrane-potential responses to current steps were analyzed, and four criteria were used to identify voltage responses that were the least affected by activation of voltage-dependent conductances. tau m was estimated from the slowest component (tau 0) of multiexponential fits of responses deemed passive by these criteria. RN was estimated from the slope of the linear region in the hyperpolarizing direction of the voltage-current relation. 4. It was not possible to measure purely passive membrane properties that were completely independent of membrane potential in any of the three classes of hippocampal neurons. Changing the membrane potential by constant current injection resulted in changes in RN and tau 0; subthreshold depolarization produced an increase, and hyperpolarization a decrease, in both RN and tau 0 for all three classes of hippocampal neurons. 5. Each of the three classes of hippocampal neurons also displayed a depolarizing "sag" during larger hyperpolarizing voltage transients. To evaluate the effect of the conductances underlying this sag on passive membrane properties, 2-5 mM Cs+ was added to the physiological saline. Extracellular Cs+ effectively blocked the sag in all three classes of hippocampal neurons, but the effect of Cs+ on RN, tau 0, and the voltage dependence of these parameters was unique for each class of neurons. 6. CA1 pyramidal neurons had an RN of 104 +/- 10 (SE) M omega and tau 0 of 28 +/- 2 ms at a resting potential of -64 +/- 2 mV (n = 12). RN and tau 0 were larger at more depolarized potentials in these neurons, but the addition of Cs+ to the physiological saline reversed this voltage dependence. 7. CA3 pyramidal neurons had an RN of 135 +/- 8 M omega and tau 0 of 66 +/- 4 ms at a resting potential of -64 +/- 1 mV (n = 14).(ABSTRACT TRUNCATED AT 400 WORDS)

Bistable membrane potential of the ciliate Coleps hirtus

2000 ◽  
Vol 203 (4) ◽  
pp. 757-764
Author(s):  
P. Rudberg ◽  
O. Sand
Keyword(s):  
Membrane Potential ◽  
Deep Level ◽  
Input Resistance ◽  
Membrane Properties ◽  
Resting Potential ◽  
Inward Rectification ◽  
Free Solution ◽  
Shallow Level ◽  
Swimming Pattern ◽  
Potential Level

In normal recording solution, the swimming pattern of the freshwater ciliate Coleps hirtus, belonging to the class Prostomatea, consists of alternating periods of nearly linear forward swimming and circular swimming within a small area. Current-clamp recordings were performed to elucidate the mechanism for this behaviour. No members of this class have previously been studied using electrophysiological techniques. The ciliates were maintained in culture and fed on the planctonic alga Rhodomonas minuta. The membrane potential showed spontaneous shifts between a more negative (deep) level of approximately −50 mV and a less negative (shallow) level of approximately −30 mV. The input resistance and capacitance at the more negative level were approximately 400 M capomega and 120 pF respectively. C. hirtus displayed a pronounced inward rectification, which was virtually insensitive to 1 mmol l(−1) Cs(+) and almost completely blocked by 1 mmol l(−1) Ba(2+). Depolarising current injections failed to evoke graded, regenerative Ca(2+) spikes. However, current-induced depolarisations from the more negative potential level (−50 mV) showed a pronounced shoulder during the repolarising phase. Increased current injections prolonged the shoulder, which occasionally stabilised at the shallow membrane potential (−30 mV). The membrane potential could be shifted to the deep level by brief hyperpolarising current injections. Similar biphasic membrane properties have not been reported previously in any ciliate. The bistability of the membrane potential was abolished in Ca(2+)-free solution containing Co(2+) or Mg(2+). In Ca(2+)-free solution containing 1 mmol l(−1) Ba(2+), brief depolarising current injections at the deep potential level evoked all-or-nothing action potentials with a prolonged plateau coinciding with the shallow potential. We conclude that the deep membrane potential in C. hirtus corresponds to the traditional resting potential, whereas the shallow level is a Ca(2+)-dependent plateau potential. In normal solution, the direction of the ciliary beat was backwards at the deep potential level and forwards at the shallow membrane potential, probably reflecting the two main phases of the swimming pattern.

Characterization of the properties of canine colonic smooth muscle in culture

1993 ◽  
Vol 265 (5) ◽  
pp. C1433-C1442 ◽  
Author(s):  
M. J. Rogers ◽  
S. M. Ward ◽  
M. A. Horner ◽  
K. M. Sanders ◽  
B. Horowitz
Keyword(s):  
Smooth Muscle ◽  
Smooth Muscles ◽  
Potential Gradient ◽  
Circular Muscle ◽  
Electrical Field Stimulation ◽  
Muscle Layer ◽  
Enteric Neurons ◽  
Resting Potential ◽  
Northern Analysis ◽  
Functional Components

We have developed and characterized an organ culture system that maintains the viability of colonic smooth muscles. Morphological, mechanical, electrical, and molecular properties of cultured canine colonic circular muscles were determined. Strips of circular muscle were cultured for up to 6 days. The smooth muscle phenotype was retained during culture; muscles contracted to agonists and responded to electrical field stimulation, suggesting that intrinsic nerves also survived in culture. Morphological analysis showed identifiable smooth muscle cells, enteric neurons, and interstitial cells, but some alterations in ultrastructure were also observed. Mechanical responses to acetylcholine suggested that the muscles developed supersensitivity during the culture period. The resting membrane potentials of cells near the submucosal surface of the circular muscle layer decreased from -82 mV on day 0 to -55 mV on day 3. Similar changes in the resting potential gradient occur when colonic muscles are treated with inhibitors of the Na(+)-K(+)-ATPase. Resting potentials of day 3 muscles remained constant in low external K+ (0.1 mM), suggesting little contribution of the pump to resting potential. Northern analysis of RNA from muscles cultured up to 6 days showed that the alpha 2-isoform of the pump decreased. The data suggest that organ-cultured strips of smooth muscle may provide a useful tool for evaluating electrical and mechanical events in conjunction with molecular analysis of functional components.

Studies of solitary semicircular canal hair cells in the adult pigeon. I. Frequency- and time-domain analysis of active and passive membrane properties

1989 ◽  
Vol 62 (4) ◽  
pp. 924-934 ◽  
Author(s):  
M. J. Correia ◽  
B. N. Christensen ◽  
L. E. Moore ◽  
D. G. Lang
Keyword(s):  
Membrane Potential ◽  
Frequency Domain ◽  
Hair Cells ◽  
Time Domain ◽  
Input Resistance ◽  
Membrane Properties ◽  
Resting Potential ◽  
The Mean ◽  
Passive Membrane Properties ◽  
Passive Membrane

1. Hair cells were enzymatically dissociated from the neuroepithelium (cristae ampullares) of the semicircular canals of white king pigeons (Columba livia). Those hair cells determined to be type II by an anatomic criterion, the ratio of the minimum width of the neck to the width of the cuticular plate, were studied with the use of the whole cell patch-clamp technique. 2. The mean +/- SD zero-current membrane potential, Vz, was found to be -54 +/- 12 mV for anterior crista hair cells (n = 71), -62 +/- 14 mV for posterior crista hair cells (n = 14), and -55 +/- 12 mV for lateral (horizontal) crista hair cells (n = 18). The mean +/- SD value of Vz for hair cells from all cristae (n = 103) was -56 +/- 13 mV. 3. Active and passive membrane properties were calculated in the time domain, in voltage- or current-clamp mode, from responses to voltage or current pulses and, in the frequency domain, by fitting a membrane model to admittance magnitude and phase data resulting from current responses to sum-of-sines voltages at different d.c. levels of voltage-clamp membrane potential. 4. The average value +/- SE of input resistance (Rin), over the range from -100 to -60 mV, was found to 1.5 +/- 0.3 G omega from a mean-voltage-as-a-function-of-current plot, V-I, (n = 7) and a mean of 1.4 +/- 0.3 G omega from individual (n = 15) current-as-a-function-of-voltage plots, I-V. A lower mean value 0.8 +/- 0.4 G omega was obtained for the input resistance from frequency-domain calculations for a different set of cells (n = 21). Also, in two different sets of cells, average input capacitance (Cin) was determined to be 12 +/- 3 pF (n = 7) from time-domain estimates and 14 +/- 3 pF (n = 21) from frequency-domain estimates. The (Rin)(Cin) product was 11 ms based on frequency-domain estimates and 17 ms from time-domain estimates. 5. I-V curves for hair cells voltage clamped at -60 mV showed some anomalous rectification for hyperpolarizations between -60 and -120 mV but no detectable N-shape for depolarizations between -50 and 90 mV. The I-V relation showed increasing slope with depolarization through the resting potential (Vz) and increased linearly between -40 and 80 mV; the best-fit straight-line maximum slope conductance for six cells over this range was 17.4 +/- 0.3 nS.(ABSTRACT TRUNCATED AT 400 WORDS)

Properties of visceral primary afferent neurons in the nodose ganglion of the rabbit

1985 ◽  
Vol 54 (2) ◽  
pp. 245-260 ◽  
Author(s):  
C. E. Stansfeld ◽  
D. I. Wallis
Keyword(s):  
Action Potentials ◽  
Input Resistance ◽  
Nodose Ganglion ◽  
Time Dependent ◽  
Membrane Properties ◽  
Resting Potential ◽  
Inward Rectification ◽  
C Cells ◽  
Axonal Conduction ◽  
A Cells

The active and passive membrane properties of rabbit nodose ganglion cells and their responsiveness to depolarizing agents have been examined in vitro. Neurons with an axonal conduction velocity of less than 3 m/s were classified as C-cells and the remainder as A-cells. Mean axonal conduction velocities of A- and C-cells were 16.4 m/s and 0.99 m/s, respectively. A-cells had action potentials of brief duration (1.16 ms), high rate of rise (385 V/s), an overshoot of 23 mV, and relatively high spike following frequency (SFF). C-cells typically had action potentials with a "humped" configuration (duration 2.51 ms), lower rate of rise (255 V/s), an overshoot of 28.6 mV, an after potential of longer duration than A-cells, and relatively low SFF. Eight of 15 A-cells whose axons conducted at less than 10 m/s had action potentials of longer duration with a humped configuration; these were termed Ah-cells. They formed about 10% of cells whose axons conducted above 2.5 m/s. The soma action potential of A-cells was blocked by tetrodotoxin (TTX), but that of 6/11 C-cells was unaffected by TTX. Typically, A-cells showed strong delayed (outward) rectification on passage of depolarizing current through the soma membrane and time-dependent (inward) rectification on inward current passage. Input resistance was thus highly sensitive to membrane potential close to rest. In C-cells, delayed rectification was not marked, and slight time-dependent rectification occurred in only 3 of 25 cells; I/V curves were normally linear over the range: resting potential to 40 mV more negative. Data on Ah-cells were incomplete, but in our sample of eight cells time-dependent rectification was absent or mild. C-cells had a higher input resistance and a higher neuronal capacitance than A-cells. In a proportion of A-cells, RN was low at resting potential (5 M omega) but increased as the membrane was hyperpolarized by a few millivolts. A-cells were depolarized by GABA but were normally unaffected by 5-HT or DMPP. C-cells were depolarized by GABA in a similar manner to A-cells but also responded strongly to 5-HT; 53/66 gave a depolarizing response, and 3/66, a hyperpolarizing response. Of C-cells, 75% gave a depolarizing response to DMPP.(ABSTRACT TRUNCATED AT 400 WORDS)

Electrogenic Na+ Transport in a Crustacean Coxal Receptor

1979 ◽  
Vol 78 (1) ◽  
pp. 29-45
Author(s):  
MAURIZIO MIROLLI
Keyword(s):  
Steady State ◽  
Membrane Potential ◽  
Input Resistance ◽  
Scylla Serrata ◽  
Partial Action ◽  
State Response ◽  
Electrogenic Na Pump ◽  
K Concentration ◽  
Low K ◽  
In Cells

1. The response of the coxal receptors of the crab Scylla serrata to step stretches consisted of a partial action potential, Vα, followed by a steady-state depolarization, V8. The input resistance of the fibre was reduced during V8. 2. In the absence of stimulation, the dendrites of the receptors depolarized when external Na+ was substituted with choline or Li+, and when the external K+ concentration was increased or decreased. The dendrites also depolarized when ouabain was added to the saline. 3. The amplitude of both Vα and V8 was dependent on external Na+. In cells which were depolarized by ouabain, the amplitude of V8 increased when the K+ concentration of the saline was reduced. 4. V8 was followed by a small, but long-lasting, after-potential which was depolarizing when the membrane potential was between −70 and −60 mV. In cells depolarized by ouabain or by low K+ saline, the after-potential became hyperpolarizing. 5. When trains of brief stretches (each 5 ms in duration) were used as stimuli, the cells responded with trains of Vα responses. During this tetanic stimulation the cells hyperpolarized; cessation of the stimulus train was followed by a long-lasting hyperpolarization (PTH). 6. PTH was abolished in Li+ saline, in low K+ saline, and in the presence of ouabain. In control or in low K+ saline, PTH was not accompanied by a decrease in the input resistance of the fibres. 7. It is concluded that an electrogenic Na+ pump (or equivalent process) contributes a substantial fraction of the membrane potential of the unstimulated coxal receptors. Pump activity could be increased by Na+-loading the distal part of the cells with trains of Vα responses. By contrast, during the steady-state response to stretch, the pump was not activated.

Beta-adrenergic actions on membrane electrical properties of dissociated smooth muscle cells

1988 ◽  
Vol 254 (3) ◽  
pp. C423-C431 ◽  
Author(s):  
H. Yamaguchi ◽  
T. W. Honeyman ◽  
F. S. Fay
Keyword(s):  
Smooth Muscle ◽  
Membrane Potential ◽  
Electrical Properties ◽  
Smooth Muscle Cells ◽  
Input Resistance ◽  
Muscle Cells ◽  
Resting Potential ◽  
Equilibrium Potential ◽  
Dependent Manner ◽  
Beta Adrenergic

Studies were carried out to determine the effects of the beta-adrenergic agent, isoproterenol (ISO), on membrane electrical properties in single smooth muscle cells enzymatically dispersed from toad stomach. In cells bathed in buffer of physiological composition, the average resting potential was -56.4 +/- 1.4 mV (mean +/- SE, n = 35). The dominant effect of exposure to ISO was hyperpolarization. The hyperpolarization was apparent in all cells studied and averaged 11.6 +/- 1.2 mV (n = 27). In the majority of the cells, hyperpolarization was accompanied by a decreased input resistance (Rin). Often the change in resistance appeared to lag behind the change in membrane potential. The lack of coincident changes in membrane potential and resistance may reflect a superposition of the outward rectification properties of the membrane on beta-adrenergic-induced increases in ionic conductance. In about half of the cells, an initial small depolarization (3.1 +/- 0.3 mV, n = 14) was accompanied by a small but distinct increase in Rin (12 +/- 2.5%). When membrane potential was made more negative than the estimated equilibrium potential for K+ (EK) by injection of current, ISO also produced biphasic effects, an initial hyperpolarization which reversed to a sustained depolarization to a value (-90 mV) near the estimated EK. The hyperpolarization by ISO could be diminished in a time-dependent manner by previous exposure to ouabain. The inhibition by ouabain, however, appeared to be a fortuitous result of glycoside-induced positive shifts in EK. These observations indicate that the dominant electrophysiological effect of beta-adrenergic stimuli is to hyperpolarize the cell membrane.(ABSTRACT TRUNCATED AT 250 WORDS)

Hyperpolarization-Activated Inward Current in Neurons of the Rat's Dorsal Nucleus of the Lateral Lemniscus In Vitro

1997 ◽  
Vol 78 (5) ◽  
pp. 2235-2245 ◽  
Author(s):  
Xiao Wen Fu ◽  
Borys L. Brezden ◽  
Shu Hui Wu
Keyword(s):  
Membrane Potential ◽  
Input Resistance ◽  
Resting Potential ◽  
Extracellular Potassium ◽  
Current Clamp ◽  
Lateral Lemniscus ◽  
Inward Current ◽  
Dorsal Nucleus ◽  
Maximal Activation

Fu, Xiao Wen, Borys L. Brezden, and Shu Hui Wu. Hyperpolarization-activated inward current in neurons of the rat's dorsal nucleus of the lateral lemniscus in vitro. J. Neurophysiol. 78: 2235–2245, 1997. The hyperpolarization-activated current ( I h) underlying inward rectification in neurons of the rat's dorsal nucleus of the lateral lemniscus (DNLL) was investigated using whole cell patch-clamp techniques. Patch recordings were made from DNLL neurons of young rats (21–30 days old) in 400 μm tissue slices. Under current clamp, injection of negative current produced a graded hyperpolarization of the cell membrane, often with a gradual sag in the membrane potential toward the resting value. The rate and magnitude of the sag depended on the amount of hyperpolarizing current. Larger current resulted in a larger and faster decay of the voltage. Under voltage clamp, hyperpolarizing voltage steps elicited a slowly activating inward current that was presumably responsible for the sag observed in the voltage response to a steady hyperpolarizing current recorded under current clamp. Activation of the inward current ( I h) was voltage and time dependent. The current just was seen at a membrane potential of −70 mV and was activated fully at −140 mV. The voltage value of half-maximal activation of I h was −78.0 ± 6.0 (SE) mV. The rate of I h activation was best approximated by a single exponential function with a time constant that was voltage dependent, ranging from 276 ± 27 ms at −100 mV to 186 ± 11 ms at −140 mV. Reversal potential ( E h) of I h current was more positive than the resting potential. Raising the extracellular potassium concentration shifted E h to a more depolarized value, whereas lowering the extracellular sodium concentration shifted E h in a more negative direction. I h was sensitive to extracellular cesium but relatively insensitive to extracellular barium. The current amplitude near maximal-activation (about −140 mV) was reduced to 40% of control by 1 mM cesium but was reduced to only 71% of control by 2 mM barium. When the membrane potential was near the resting potential (about −60 mV), cesium had no effect on the membrane potential, current-evoked firing rate and input resistance but reduced the spontaneous firing. When the membrane potential was more negative than −70 mV, cesium hyperpolarized the cell, decreased current-evoked firing and increased the input resistance. I h in DNLL neurons does not contribute to the normal resting potential but may enhance the extent of excitation, thereby making the DNLL a consistently powerful inhibitory source to upper levels of the auditory system.

Input Resistance, Time Constants, and Spike Initiation

1998 ◽  
Author(s):  
Christof Koch
Keyword(s):  
Membrane Potential ◽  
Time Constant ◽  
Action Potentials ◽  
Input Resistance ◽  
Current Injection ◽  
Membrane Conductances ◽  
Voltage Dependent ◽  
Dc Component ◽  
Definition Of ◽  
A Current

This chapter represents somewhat of a tephnical interlude. Having introduced the reader to both simplified and more complex compartmental single neuron models, we need to revisit terrain with which we are already somewhat familiar. In the following pages we reevaluate two important concepts we defined in the first few chapters: the somatic input resistance and the neuronal time constant. For passive systems, both are simple enough variables: Rin is the change in somatic membrane potential in response to a small sustained current injection divided by the amplitude of the current injection, while τm is the slowest time constant associated with the exponential charging or discharging of the neuronal membrane in response to a current pulse or step. However, because neurons express nonstationary and nonlinear membrane conductances, the measurement and interpretation of these two variables in active structures is not as straightforward as before. Having obtained a more sophisticated understanding of these issues, we will turn toward the question of the existence of a current, voltage, or charge threshold at which a biophysical faithful model of a cell triggers action potentials. We conclude with recent work that suggests how concepts from the subthreshold domain, like the input resistance or the average membrane potential, could be extended to the case in which the cell is discharging a stream of action potentials. This chapter is mainly for the cognoscendi or for those of us that need to make sense of experimental data by comparing therp to theoretical models that usually fail to reflect reality adequately. In Sec. 3.4, we defined Kii (f) for passive cable structures as the voltage change at location i in response to a sinusoidal current injection of frequency f at the same location. Its dc component is also referred to as input resistance or Rin. Three difficulties render this definition of input resistance problematic in real cells: (1) most membranes, in particular at the soma, show voltage-dependent nonlinearities, (2) the associated ionic membrane conductances are time dependent and (3) instrumental aspects, such as the effect of the impedance of the recording electrode on Rin, add uncertainty to the measuring process.

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