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**Solutions to Homework Assignment #2
**

1. The results of a voltage-clamp experiment are shown below.

−30

1

−40

0.9

−45

0.8

0.7

−50

−50

0.6

0

50

100

Time (ms)

150

200

0.5

y∞(V)

V (mV)

−35

0.4

0

0.3

0.2

−7

I (nA)

0.1

−14

−70

−60

−50

−40

V (mV)

−30

0

−20

−21

−50

0

50

100

Time (ms)

150

200

Assume that the measured transmembrane current I (V , t ) is comprised of a passive leakage

**current I L (V ) = g L (V − E L ) and a time- and voltage-dependent current I y (V , t ) = g y y (V − E y ) ,
**

where the dynamics of the gating particle y are first-order, i.e., dy (V , t ) dt = ( y∞ − y ) τ y .

The time-constant τ y is independent of voltage, whereas the voltage dependence of the

steady-state value y∞ (V ) is shown in the figure above.

It is known that the Nernst

**equilibrium potential E y = −70 mV .
**

a. Is the current I y (V , t ) activated by depolarization or hyperpolarization? Is I y (V , t ) an

inward or an outward current in the voltage-clamp experiment performed above?

b. From the results of the voltage-clamp experiment, find the values of g L , E L and g y .

(25 pts)

a. The asymptotic value of the activation particle y increases if the transmembrane potential is

made more positive, so the current I y (V , t ) is activated by depolarization. In the voltageclamp experiment, hyperpolarization of the membrane leads to a larger total inward current.

Because hyperpolarization causes deactivation of the current I y (V , t ) , it follows that this must

be an outward current. This can also be argued from the value of the equilibrium potential –

because E y is more negative than the holding potentials used in the voltage-clamp experiment,

the term V − E y must be positive, as are g y and y , so I y (V , t ) must be a positive, i.e., outward,

current.

Dr. Ian C. Bruce

March 8, 2005

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and therefore: −21×10−9 = g L ( −50 × 10−3 − EL ) + g y ⋅ 0.3 ⋅ ( −50 × 10−3 + 70 × 10−3 ) . so we must construct at least three simultaneous equations from the voltage-clamp results. I = 0 and y = 1 . At t = 100 ms . Dr. There are three unknowns. For t < 0 . At t ≈ 99 ms . I = −21×10−9 and y = 0. Solving any three of these equations simultaneously gives: g L = 200 nS. These are based on the ionic current equation: I = I L + I y = g L (V − EL ) + g y y (V − E y ) evaluated for at least three different time points in the experiment.3 . and therefore: −14 ×10−9 = g L ( −30 ×10−3 − EL ) + g y ⋅ 0. I = −14 ×10−9 and y = 1 .b. and therefore: −14 ×10−9 = g L ( −50 × 10−3 − EL ) + g y ⋅1⋅ ( −50 × 10−3 + 70 × 10−3 ) . 2005 Page 2 of 6 .3 . At t = 0 .3 ⋅ ( −30 ×10−3 + 70 ×10−3 ) . and therefore: 0 = g L ( −30 ×10−3 − EL ) + g y ⋅1⋅ ( −30 ×10−3 + 70 × 10−3 ) . Bruce March 8. I = −14 ×10−9 and y = 0. Ian C. g y = 500 nS. EL = +70 mV.

2. re ≈ 0 ). Assuming that the axons can be approximated by infinite cables: a.2 = Z 0. π 2d 3 and consequently: Z 0.1 ( d1 ) ( d2 ) (10) −3 2 32 32 c. 4 Ri and consequently: d2 λ2 = = 1. What are the relative input resistances of the two cables? c. What are the relative time constants of the two cables? (20 pts) a. Consequently.1ri . The time constant: τ = rm cm = Rm ⋅ C π d = RmCm πd m is independent of the fiber diameter. The input resistance of a semi-infinite cable is rm ri . so the time constant τ 2 of cable 2 relative to the time constant τ 1 of cable 1 is τ 2 τ 1 = 1 . 2005 Page 3 of 6 .1 .3536 . For a cylindrical cable of diameter d : rm ri = Rm 4 Ri ⋅ = πd πd2 4 Rm Ri .e. Dr. the conduction velocity θ 2 for cable 2 ( d 2 = 10 μm ) relative to the conduction velocity θ1 for cable 1 ( d1 = 5 μm ) is: d2 θ2 10 = = = 1. the space constant is: λ= d Rm . The input resistance of an infinite cable is half of this.2 rm . Bruce March 8.1 rm . For an unmyelinated axon of diameter d . You have two unmyelinated axons.4142 . λ1 d1 d. the propagation velocity is proportional to d . so the relative input resistances are the same as for a semi-infinite cable: Z 0. What are the relative conduction velocities of the two cables? b.2 ( d 2 ) (d ) ( 5) = 0. θ1 d1 5 b. = = 1 32 = 32 −3 2 Z 0.4142 . identical except that their diameters are 5 μm and 10 μm. Assuming a negligible extracellular resistance (i. What are the relative space constants of the two cables? d.2 ri . Ian C..

If the late outward current is carried by potassium and EK ≈ −65 mV for this axon. e. d. Because this current turns on instantaneously and does not vary with time. What is the approximate threshold potential Vthr of this axon? e. What caused the small sustained inward current when the membrane was stepped to Vc = −125 mV ? b. a. so Vthr ≈ −35 mV . The voltage-gated currents turn on somewhere between −40 and −30 mV. Why did the peak amplitude of the early inward current decrease when Vc was increased from ~ 0 mV to +40 mV ? d. The results can be interpreted by considering a parallelconductance model with voltagegated sodium and potassium channels and a passive leakage channel. and consequently no potassium current reversals are observed. Why did the peak amplitude of the early inward current ( at t ≈ 1 ms ) increase when Vc was increased from −30 mV to ~ 0 mV ? c. c. Over this range of voltages. why is there no reversal of the late outward current in any of these measurements? (30 pts) a. This is produced by activation of this ion channel → more open channels → increased conductance → increased current. The membrane was initially held at voltage Vh = −60 mV and was stepped at time t = 0 to voltage Vc shown to the right of each current trace. so E Na ≈ +50 mV . Transmembrane currents measured in a squid axon from a voltage-clamp experiment are shown below. but the membrane potential approaches the ion’s Nernst equilibrium potential → decreased current. what is the approximate sodium equilibrium potential ENa for this axon? f. it must be the leakage current in the HH model. The potassium current activates somewhere above −30 mV.3. f. so the potassium channels are all closed at the potassium reversal potential of EK ≈ −65 mV . Ian C. The early current reverses somewhere between +40 and +60 mV. Dr. b. the conductance does not change much. 2005 Page 4 of 6 . If the early inward current is carried by sodium. Bruce March 8.

Bruce I0 2 z 2 − h2 4πσ e ( h 2 + z 2 )5 2 .e. the extracellular potential as a function of z . ∂ 2φe is then: ∂z 2 ∂φe I z =− 0 ∂z 4πσ e ( h 2 + z 2 )3 2 ⇒ 2 2 2 I 0 3z − ( h + z ) ∂ 2φe = ∂z 2 4πσ e ( h 2 + z 2 )5 2 = Dr. h is the distance between the source and the fiber where the source is perpendicular to the fiber. i.. March 8. i.e. b. 2005 Page 5 of 6 .5 of Plonsey and Barr (shown on p. (25 pts) a. φe ( z ) . The activation function .70): φe = I0 . ∂z 2 c. Therefore the extracellular potential at position z on the fiber is: φe ( z ) = I0 4πσ e 1 h + z2 2 b. Find: a. the activation function ∂ 2φe .4. 8 of Lecture #19). (7.. 7. the positions of the boundaries between the initial depolarized and hyperpolarized regions. stating which are a peak of depolarization and which are peaks of hyperpolarization. 4πσ e r where r is the distance from the source to the fiber at the axial distance z from the middle of the fiber. at z = 0 . Consider the source-fiber geometry of Fig. such that the extracellular potential is given by Eqn. the positions of the peaks of the resulting regions of initial depolarization and hyperpolarization predicted by the activation function. Ian C. The distance r from the electrode to the position z on the fiber is: r = h2 + z2 . and d.

4 2 ∂ φe/∂z (normalized) 0 2 −0. The boundaries between the depolarized and hyperpolarized regions are located where the activation function equals zero: ∂ 2φe I0 2 z 2 − h2 = =0 ∂z 2 4πσ e ( h 2 + z 2 )5 2 ⇒ 2 z 2 − h2 ( h2 + z 2 ) 52 =0 ⇒ 2z2 − h2 = 0 ⇒z=± 1 2 h. ± 3 2 h. 2005 5 Page 6 of 6 .c. Bruce −√3/2 −√1/2 0 √1/2 √3/2 z March 8. z = 0 is the location of the peak of the hyperpolarized region.2 −0. The peaks of the activation function ∂ 2φe are located where its derivate equals zero: ∂z 2 ∂ 3φe I 0 9 zh 2 − 6 z 3 = =0 ∂z 3 4πσ e ( h 2 + z 2 )7 2 ⇒ 9 zh 2 − 6 z 3 (h 2 + z2 ) 72 =0 ⇒ 9 zh 2 − 6 z 3 = 0 ⇒ z 3 = 23 zh 2 ⇒ z = 0. h=1 0. and z = ± 3 2 h are the locations of the peaks of the flanking depolarized regions.8 −1 −5 Dr. Ian C.6 −0. d.2 −0.

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