Stimuli and Local Response
For a cell to become excited, it must be acted upon by a stimulus. Stimuli are classified as subthreshold, threshold, and suprathreshold.
With a weak (subthreshold) stimulus, the resting potential decreases only slightly at the electrode site, producing a local response (LR). Its main feature is a direct proportionality: the stronger the stimulus, the higher the response amplitude.
If the stimulus reaches the threshold value, the local response reaches the critical firing level (threshold of depolarization). At this exact moment, the local membrane change transitions into a full-fledged action potential.
Dynamics and Phases of the Action Potential
The development of excitation involves several consecutive stages displayed on a graph:
- Pre-spike (Depolarization prepotential): A slow partial depolarization corresponding to the local response.
- Depolarization (Spike, peak): A rapid voltage surge. The potential shifts from the critical threshold (-50 mV) to positive values (e.g., +30 mV). This polarity reversal is called overshoot (the inner surface of the membrane becomes positive).
- Repolarization: Voltage drops and returns toward the resting level. At the end of this phase, a negative after-potential (NAP) or after-depolarization may occur.
- After-hyperpolarization: The potential drops below the baseline -70 mV. This is designated as the positive after-potential (PAP).
The duration and shape of these phases depend on the tissue type. In neurons and skeletal myocytes, the spike is very brief, and after-potentials are variable.
Ionic Mechanisms and Channel Function
All potential changes are driven by transmembrane tubular proteins known as ion channels.
- Resting state: The membrane potential is -70 mV. Potassium channels are closed. Sodium channels are also closed but primed for activation: their activation m-gates are shut, and inactivation h-gates are open. Potassium concentration is higher inside the cell, while sodium is higher outside.
- Onset of stimulation (LR): The m-gates begin to slowly open, but the inward sodium current remains negligible.
- Depolarization phase: Upon reaching the threshold (-50 mV), all m-gates open. An avalanche-like, passive influx of Na⁺ ions begins down their concentration gradient.
- Repolarization: The incoming positive charge triggers the closure of the h-gates (inactivation of sodium channels). Simultaneously, voltage sensors on potassium pores respond by opening. K⁺ ions rush outward, restoring the negative charge inside.
- Hyperpolarization and restoration: Because potassium channels close slowly, K⁺ efflux continues (sometimes supplemented by Cl⁻ influx), driving the potential below -70 mV. Finally, the Na⁺/K⁺-ATPase actively pumps ions back to baseline. Sodium channels reset (closing m-gates and opening h-gates), and the cell is ready for a new impulse.
The All-or-None Law
This law reflects the constancy of the action potential amplitude. Its core principle is simple:
- A subthreshold stimulus produces no impulse ("none").
- A threshold or suprathreshold stimulus generates an AP of the maximum possible amplitude for that specific cell ("all").
Single structures obey this law: neurons, individual nerve and muscle fibers, smooth muscle, and cardiac tissue. However, whole nerve trunks or skeletal muscles do not obey it because they consist of numerous fibers with varying excitation thresholds.