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At point C in Figure 5-18, the stimulus is even stronger. Now the local potential has barely reached the level required to elicit an action potential, called the threshold level, but this occurs only after a short "latent period." At point D, the stimulus is still stronger, the acute local potential is also stronger, and the action potential occurs after less of a latent period.

Thus, this figure shows that even a very weak stimulus causes a local potential change at the membrane, but the intensity of the local potential must rise to a threshold level before the action potential is set off.

"Refractory Period" After an Action Potential, During Which a New Stimulus Cannot Be Elicited

A new action potential cannot occur in an excitable fiber as long as the membrane is still depolarized from the preceding action potential. The reason for this is that shortly after the action potential is initiated, the sodium channels (or calcium channels, or both) become inactivated, and no amount of excitatory signal applied to these channels at this point will open the inactivation gates. The only condition that will allow them to reopen is for the membrane potential to return to or near the original resting membrane potential level. Then, within another small fraction of a second, the inactivation gates of the channels open, and a new action potential can be initiated.

The period during which a second action potential cannot be elicited, even with a strong stimulus, is called the absolute refractory period. This period for large myelinated nerve fibers is about 1/2500 second. Therefore, one can readily calculate that such a fiber can transmit a maximum of about 2500 impulses per second.

Inhibition of Excitability— "Stabilizers" and Local Anesthetics

In contrast to the factors that increase nerve excitability, still others, called membrane-stabilizing factors, can decrease excitability. For instance, a high extracellular fluid calcium ion concentration decreases membrane permeability to sodium ions and simultaneously reduces excitability. Therefore, calcium ions are said to be a "stabilizer."

Local Anesthetics. Among the most important stabilizers are the many substances used clinically as local anesthetics, including procaine and tetracaine. Most of these act directly on the activation gates of the sodium channels, making it much more difficult for these gates to open, thereby reducing membrane excitability. When excitability has been reduced so low that the ratio of action potential strength to excitability threshold (called the "safety factor") is reduced below 1.0, nerve impulses fail to pass along the anesthetized nerves.

Recording Membrane Potentials and Action Potentials

Cathode Ray Oscilloscope. Earlier in this chapter, we noted that the membrane potential changes extremely rapidly during the course of an action potential. Indeed, most of the action potential complex of large nerve fibers takes place in less than 1/1000 second. In some figures of this chapter, an electrical meter has been shown recording these potential changes. However, it must be understood that any meter capable of recording most action potentials must be capable of responding extremely rapidly. For practical purposes, the only common type of meter that is capable of responding accurately to the rapid membrane potential changes is the cathode ray oscilloscope.

Figure 5-19 shows the basic components of a cathode ray oscilloscope. The cathode ray tube itself is composed basically of an electron gun and a fluorescent screen against which electrons are fired. Where the electrons hit the screen surface, the fluorescent material glows. If the electron beam is moved across the screen, the spot

Recorded action potential Horizontal

Recorded action potential Horizontal

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Figure 5-19

Cathode ray oscilloscope for recording transient action potentials.

of glowing light also moves and draws a fluorescent line on the screen.

In addition to the electron gun and fluorescent surface, the cathode ray tube is provided with two sets of electrically charged plates—one set positioned on the two sides of the electron beam, and the other set positioned above and below. Appropriate electronic control circuits change the voltage on these plates so that the electron beam can be bent up or down in response to electrical signals coming from recording electrodes on nerves. The beam of electrons also is swept horizontally across the screen at a constant time rate by an internal electronic circuit of the oscilloscope. This gives the record shown on the face of the cathode ray tube in the figure, giving a time base horizontally and voltage changes from the nerve electrodes shown vertically. Note at the left end of the record a small stimulus artifact caused by the electrical stimulus used to elicit the nerve action potential. Then further to the right is the recorded action potential itself.

References

Alberts B, Johnson A, Lewis J, et al: Molecular Biology of the Cell. New York: Garland Science, 2002. Grillner S: The motor infrastructure: from ion channels to neuronal networks. Nat Rev Neurosci 4:573, 2003.

Hodgkin AL: The Conduction of the Nervous Impulse. Springfield, IL: Charles C Thomas, 1963.

Hodgkin AL, Huxley AF: Quantitative description of membrane current and its application to conduction and excitation in nerve. J Physiol (Lond) 117:500,1952.

Kleber AG, Rudy Y: Basic mechanisms of cardiac impulse propagation and associated arrhythmias. Physiol Rev 84:431, 2004.

Lu Z: Mechanism of rectification in inward-rectifier K+ channels. Annu Rev Physiol 66:103, 2004.

Matthews GG: Cellular Physiology of Nerve and Muscle. Malden, MA: Blackwell Science, 1998.

Perez-Reyes E: Molecular physiology of low-voltage-activated T-type calcium channels. Physiol Rev 83:117, 2003.

Poliak S, Peles E: The local differentiation of myelinated axons at nodes of Ranvier. Nat Rev Neurosci 12:968,2003.

Pollard TD, Earnshaw WC: Cell Biology. Philadelphia: Elsevier Science, 2002.

Ruff RL: Neurophysiology of the neuromuscular junction: overview. Ann N Y Acad Sci 998:1, 2003.

Xu-Friedman MA, Regehr WG: Structural contributions to short-term synaptic plasticity. Physiol Rev 84:69, 2004.

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