Article

# Effects of external currents on duration and amplitude of normal and prolonged action potentials from single nodes of Ranvier

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## Abstract

Duration and amplitude of normal and prolonged action potentials from single nodes of Ranvier vary as functions of potential changes induced by currents from an external source. The quantitative relations between externally applied potential and the resulting potential generated within the system are analyzed in order to obtain information about the kinetics of the electromotance,—potential,—and chemical changes taking place during excitation. The following preliminary conclusions are drawn: A depolarizing and a repolarizing process (positive and negative electromotance) increase and decrease with the potential. For a sudden potential displacement the negative electromotance reaches its new value at a faster rate than the positive electromotance. Since the individual values of the two electromotances depend on the potential and since they both generate a potential which is proportional to the difference of their absolute values, the values of either electromotance are determined by this difference as well as by any externally induced potential change.

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... The effect of maintained polarization upon the physiological properties of the node in potassium-rich media has been reported recently by Mueller (1958a) and by Stiimpfli (1958). They observed discontinuous variations in the membrane potential when the strength of the anodal polarizing current through the node was gradually increased. ...
... However, this sort of action potential has been reported to be elicited only under abnormal conditions. For instance, the hyperpolarizing response can be elicited only on prior depolarization of squid (Segal, 1958;Tasaki, 1959a), frog (Mueller, 1958;Stampfli, 1959;Liittgau, 1960), or toad (Tasaki, 1959a) axons. Although quantitative data on the kinetics of their electrogenetic processes are not as yet available, Grundfest (1960) suggested that these responses may develop as a result of reinstated UK-inactivation" when the K-conductance is first increased under experimental conditions. ...
Article
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Article
THE electrical properties of the myelinated nerve fibre in potassium-rich solution were observed by Stämpni1, Mueller2 and Tasaki3. Those of the unmyelinated nerve fibre were observed by Segal4, Tasaki3 and Moore5. When the membrane potential is depolarized by potassium, the potential difference across the membrane produced by the inward current pulse is larger than that produced by the outward one and exhibits a kind of discontinuous increase at a certain current intensity (the current-potential relation is non-linear). The time-course of the potential difference produced by the inward current has two steps. This phenomenon is called anodal threshold phenomenon'' by Segal4 and hyperpolarizing response'' by Tasaki3. Tasaki reported that the hyperpolarizing response in the giant axon of the squid was slightly different from that in the myelinated nerve fibre.
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Article
Previous step voltage-clamp measurements on frog skin showed the presence of an N-shaped current-potential (I-V) relation in excitable skin. However, the collection and reconstruction of I-V data using discrete step changes of skin potential was tedious because of the long refractory period (up to 1 min) in frog skin. A direct and rapid (5 msec) method for recording the N-shaped I-V characteristic in real time is presented. Ramp functions are used as the command to the clamp system instead of a step function. Consequently the skin potential is forced to change in a linear manner (as commanded) and the skin current can be recorded as a continuous function of the controlled change of skin potential. With the ramp clamp, a low-resistance membrane state (〈 10 Ω · cm2) resembling a breakdown phenomenon was observed at high skin potential (〉 300 mv). Entry into the low resistance state resulted in a collapse of the N-shaped I-V relation to a nearly linear function. The utility of the ramp measurement is demonstrated by predicting (1) that the maximum rate of rise of the spike occurs at a voltage corresponding to the valley (local minimum) in the N-shaped I-V curve, (2) that the rate of rise of the spike increases with increasing clamp currents, (3) the voltage peak of the spike, and (4) the time course of the rising phase of the spike.
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Article
It is shown in a mathematical model of a myelinated nerve fiber that the development of a local response in an inexcitable node plays an important role in the mechanism of the "jumping" of an action potential (AP) across the inexcitable node. In the absence of such a response (for example, in the case of a 1000-fold decrease in the maximum sodium permeability, $$\bar P$$ Na) in fibers with normal relations between the length of the internodal segment (L) and its diameter (D) (L/D>100), the conduction is blocked. It is possible only in fibers with relatively short internodal segments (L/D
Article
1. Current flow outward through the caudal, reactive membrane of the cell causes direct stimulation of the electroplaque. The electrical response in denervated as well as in normal preparations recorded with internal microelectrodes is first local and graded with the intensity of the stimulus. When membrane depolarization reaches about 40 mv. a propagated, all-or-nothing spike develops. 2. Measured with internal microelectrodes the resting potential is 73 mv. and the spike 126 mv. The latter lasts about 2 msec. and is propagated at approximately 1 M.P.S. 3. The latency of the response decreases nearly to zero with strong direct stimulation and the entire cell may be activated nearly synchronously. 4. Current flow inward through the caudal membrane of the cell does not excite the latter directly, but activation of the innervated cell takes place through stimulation of the nerve terminals. This causes a response which has a latency of not less than 1.0 msec. and up to 2.4 msec. 5. The activity evoked by indirect stimulation or by a neural volley includes a prefatory potential which has properties different from the local response. This is a postsynaptic potential since it also develops in the excitable membrane which produces the local response and spike. 6. On stimulation of a nerve trunk the postsynaptic potential is produced everywhere in the caudal membrane, but is largest at the outer (skin) end of the cell. The spike is initiated in this region and is propagated at a slightly higher rate than is the directly elicited response. Strong neural stimulation can excite the entire cell to simultaneous discharge. 7. The postsynaptic potential caused by neural or indirect stimulation may be elicited while the cell is absolutely refractory to direct excitation. 8. The postsynaptic potential is not depressed by anodal, or enhanced by cathodal polarization. 9. It is therefore concluded that the postsynaptic potential represents a membrane response which is not electrically excitable. Neural activation of this therefore probably involves a chemical transmitter. 10. The nature of the transmitter is discussed and it is concluded that this is not closely related to acetylcholine. 11. Paired homosynaptic excitation discloses facilitation which is not present when the conditioning stimulus is direct or through a different nerve trunk. These results may be interpreted in the light of the existence of a neurally caused chemical transmitter or alternatively as due to presynaptic potentiation. 12. The electrically excitable system of the electroplaque has two components. In the normal cell a graded reaction of the membrane develops with increasing strength of stimulation until a critical level of depolarization, which is about 40 mv. 13. At this stage a regenerative explosive reaction of the membrane takes place which produces the all-or-nothing spike and propagation. 14. During early relative refractoriness or after poisoning with some drugs (eserine, etc.) the regenerative process is lost. The membrane response then may continue as a graded process, increasing proportionally to the stimulus strength. Although this pathway is capable of producing the full membrane potential the response is not propagated. 15. Propagation returns when the cell recovers its regenerative reaction and the all-or-nothing response is elicited. 16. Excitable tissues may be classified into three categories. The axon is everywhere electrically excitable. The skeletal muscle fiber is electrically excitable everywhere except at a restricted region (the end plate) which is only neurally or chemically excitable. The electroplaque of the eel, and probably also cells of the nervous system have neurally and electrically excitable membrane components intermingled. The electroplaques of Raia and probably also of Torpedo as well as frog muscle fibers of the "slow" system have membranes which are primarily neurally and chemically excitable. Existence of a category of invertebrate muscle fibers with graded electrical excitability is also considered. 17. In the eel electroplaque and also probably in the cells of neurons, tests of the mode of neural activation carried out by direct or antidromic stimulation cannot reveal the neurally and chemically activated component. The data of such tests though they appear to prove electrical transmission are therefore inadequate for the detection and study of the chemically initiated process.
Article
The duration of action potentials from single nodes of Ranvier can be increased by several methods. Extraction of water from the node (e.g. by 2 to 3 M glycerin) causes increased durations up to 1000 msec. 1 to 5 min. after application of the glycerin the duration of the action potential again decreases to the normal value. Another type of prolonged action potential can be observed in solutions which contain K or Rb ions at concentrations between 50 mM and 2 M. The nodes respond only if the resting potential is restored by anodal current. The kinetics of these action potentials is slightly different. Their maximal durations are longer (up to 10 sec.). Like the normal action potential, they are initiated by cathodal make or anodal break. They also occur in external solutions which contain no sodium. The same type of action potentials as in KCl is found when the node is depolarized for some time (15 to 90 sec., 100 to 200 mv.) and is then stimulated by cathodal current. These action potentials require no K or Na ions in the external medium. Their maximal duration increases with the strength and duration of the preceding depolarization. The possible origin of the action potentials in KCl and after depolarization, and their relation to the normal action potentials and the negative after-potential are discussed.
Ueber die Ausloesung rhythmischer und nichtrhythmischer Er-regtmgen im peripheren Nerven
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