Ion channel

An ion channel is a membrane protein that provides a selective pathway for ions across a cell membrane. An open pore permits passive transport down the relevant electrochemical gradient; it does not supply the free energy for uphill transport as a pump involved in active transport does. Two things are needed for a net electric current: an available pathway and an electrochemical potential difference. Opening a pore alone need not produce a net electric current. See Ion Channels and the Electrical Properties of Membranes.

The selective permeability of an ion channel depends on interactions between its pore and the passing ions, not simply on a hole's diameter. Some ion channels strongly favor potassium over sodium; others admit several kinds of positively charged ions. Thus selective does not necessarily mean exclusive to one ion species. See Purves et al., Neuroscience, 2nd edition (2001): molecular structure of ion channels and the mixed-channel example in Purves et al.: principles derived from studies of the neuromuscular junction.

Gating is the opening and closing of an ion channel's pathway. A voltage-gated ion channel responds to membrane potential; an ionotropic receptor can respond to a bound ligand. Gating changes the available electrical conductance, while the electrochemical gradient supplies the drive for passive transport. For a fixed reversal potential in the simple model below, opening more identical ion channels increases the magnitude of the electric current at a fixed nonzero drive, without choosing its sign independently of membrane potential. The two-state ion-channel gating model describes how the open fraction can change. See the Alberts reference above.

For a worked approximation, hold an open population's electrical conductance at $g=10\mathrm{nS}$ and its reversal potential at $E_{\mathrm{rev}}=-80\mathrm{mV}$. Use $V=V_{\mathrm{inside}}-V_{\mathrm{outside}}$ for membrane potential and inward-positive conventional current:At $V=-100,-80,-60\mathrm{mV}$, respectively, the electric currents are $+200,0,-200\mathrm{pA}$, since $1\mathrm{nS}\times 1\mathrm{mV}=1\mathrm{pA}$. Positive electric current means net positive electric charge entering, or net negative electric charge leaving; it does not mean that every ion moves inward. For a pore passing only positive ions, the three results mean net inward passage, no net passage, and net outward passage. These stipulated values describe an illustrative Ohmic approximation, not measured properties of a particular ion channel; see reversal potential for the current plot and limitations.

The useful connection is between a molecular gate and a controllable electric current. A ligand can change electrical conductance through an ionotropic receptor, or membrane potential can change it through a voltage-gated ion channel, connecting recognition and electrical signaling. In a historical landmark, Erwin Neher and Bert Sakmann reported currents from individual ion channels in frog muscle membranes in 1976. See Neher and Sakmann, Single-channel currents recorded from membrane of denervated frog muscle fibres, Nature 260, 799–802 (1976).

The probability that an ion channel is in a conducting open state under specified conditions. For N equivalent channels, each carrying open-state electric current i, the mean total electric current is $N{P}_{O}i$. An ensemble open fraction and the fraction of time one channel remains open coincide under suitable stationary and ergodic assumptions. A two-state ion-channel gating model is one model of this probability, not its definition.

For N independent equivalent ion channels with open probability $P_O$ and approximately Ohmic open-state electrical conductance $\gamma$, the mean electric current is $I=N{P}_O\gamma (V-E)$. A numerical Electrical conductance cannot be inferred without the relevant reversal potential or driving voltage.

An ion channel that conducts chloride and, for some family members, other anions. Members differ in voltage dependence and physiological role; an anion channel is distinct from a coupled anion exchanger.

A guard-cell outward-rectifying potassium channel. Membrane depolarization favors potassium efflux through this channel during stomatal closure.

A slow anion channel important for guard-cell anion efflux. Regulatory phosphorylation links ABA-responsive and calcium-dependent kinases to channel activation.

A mechanosensitive ion channel changes gating in response to mechanical conditions transmitted through the membrane or associated structures. A changed calcium signal alone does not identify which channel or mechanical coupling was responsible.

A large-conductance bacterial mechanosensitive channel serving as an emergency release pathway under high membrane tension.

A bacterial mechanosensitive channel that can release solutes during sudden osmotic downshock.

Channel desensitization is entry into a less responsive or nonconducting state during sustained stimulation. A ligand-bound channel can therefore be closed without the ligand having dissociated.

A ligand-gated ion channel couples ligand occupancy to the probability of pore opening. The ligand-binding site and ion pathway can be distinct, and sustained occupancy can lead to a desensitized nonconducting state.

A selectivity filter is a channel-pore region that preferentially stabilizes permeating ions through geometry and chemical interactions. Selectivity balances dehydration and coordination energies rather than following pore diameter alone.