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WIREs Membr Transp Signal

Biophysical properties of the voltage‐gated proton channel H _V 1

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Boris Musset, Thomas DeCoursey

Published Online: May 11 2012

DOI: 10.1002/wmts.55

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The biophysical properties of the voltage‐gated proton channel (H_V1) are the key elements of its physiological function. The voltage‐gated proton channel is a unique molecule that in contrast to all other ion channels is exclusively selective for protons. Alone among proton channels, it has voltage‐ and time‐dependent gating like other ‘classical’ ion channels. H_V1 is furthermore a sensor for the pH in the cell and the surrounding media. Its voltage dependence is strictly coupled to the pH gradient across the membrane. This regulation restricts opening of the channel to specific voltages at any given pH gradient, therefore allowing H_V1 to perform its physiological task in the tissue it is expressed in. For H_V1 there is no known blocker. The most potent channel inhibitor is zinc (Zn²⁺) which prevents channel opening. An additional characteristic of H_V1 is its strong temperature dependence of both gating and conductance. In contrast to single‐file water‐filled pores like the gramicidin channel, H_V1 exhibits pronounced deuterium effects and temperature effects on conduction, consistent with a different conduction mechanism than other ion channels. These properties may be explained by the recent identification of an aspartate in the pore of H_V1 that is essential to its proton selectivity. WIREs Membr Transp Signal 2012, 1:605–620. doi: 10.1002/wmts.55

Figure 1.

Proton movement in a single water file pore. The proton hops on the first water molecule and each intermediate hydronium releases a proton to the nearest oxygen in the next water molecule. The picture below shows the Newton cradle as a mnemonic device for proton movement through a single file of water.

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Figure 2.

Kinetic of outward‐ and tail currents. The figure shows the outward current fitted by a single exponential (red). There is a short delay before the exponential starts. The maximal value for the exponential gives the value for the maximal proton current at this voltage. Here the end of the pulse and the maximal current are almost the same. The tail current is perfectly describable with a single exponential (red).

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Figure 3.

End of pulse. The figure shows a current trace during a pulse protocol. The amplitude of the current at the end of the pulse (I_end) is depicted together with the amplitude of the maximal tail current (I_tail) minus the leak current. The reversal potential (V_rev) is calculated with the equation in the center of the current trace. The reversal potential thereby is able to reflect changes in pHi.

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Figure 4.

Threshold of activation in voltage‐gated proton channels. Current family shows in red the first appearance of proton channels in outward and tail current. On the right the voltage pulses are displayed exhibiting in red the threshold voltage.

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Figure 5.

Tail currents to determine the reversal potential. A prepulse to 50 mV is given followed by pulses to −30 through 20 mV. At 0 mV the tail current is nearly a straight line indicating the reversal potential (pH_i = pH_o = 7).

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Figure 6.

Enhanced gating mode g–V shift. The figure shows the conductance voltage plot (g–V) of a human monocyte before and after PMA stimulation. The ‘enhanced gating mode’ is detectable as a left shift in the g–V curve. The pH_i = pH_o = 7 in perforated patch mode.

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Figure 7.

Secondary structure of H_V1 and voltage‐gated cation channels. S1–S3 are depicted in yellow, S4 orange, and S5–S6 in red. The general concept of S1–S4 in the proton channel and voltage‐gated cation channels is the same with S4 as voltage sensor. The proton channel lacks S5–S6 the pore domain.

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