The positive and negative heat production associated with a nerve impulse

1958 ◽  
Vol 148 (931) ◽  
pp. 149-187 ◽  
Keyword(s):  
Chemical Reactions ◽  
Heat Production ◽  
Surface Membrane ◽  
Nerve Impulse ◽  
Active Phase ◽  
Fibre Surface ◽  
Excitable Membrane ◽  
Substantial Part ◽  
Recording Equipment ◽  
Muscle Twitch

The ‘initial’ heat production of a non-medullated nerve ( Maia ) has been reinvestigated with more rapid recording equipment than was previously available. In a single impulse at 0° C a positive heat production was observed averaging about 9 x 10 -6 cal/g nerve: this is rapid and is probably associated with the active phase of the impulse. It is followed by a rather slower heat absorption averaging about 7 x 10 -6 cal/g nerve and lasting for about 300 ms. Previous methods were too slow to do more than record the difference between the two, the ‘net heat’, viz. about 2 x 10 -6 cal/g nerve: this is about one-third greater at 0°C than at 18° C. Maia nerves contain fibres from 20 to 0.3 µ in diameter, and about half the heat is probably derived from fibres less than 3.0 µ . The velocities of impulses in them at 0° C vary from 1.4 to 0.1 m/s, so impulses reach the recording thermojunctions throughout a long interval. Thus the observed course of the heat production is the resultant of positive and negative components in different fibres, and a substantial part of each is masked. The real positive and negative heats, therefore, are substantially greater than those observed: on the most likely estimate of velocity distribution, in a single impulse at 0° C they are about 14 x 10 -6 cal/g and — 12 x 10 -6 cal/g, respectively. Heat production, like ionic interchange, is probably proportional to fibre surface, which in 1 g of Maia nerve is estimated as 10 4 cm 2 . If the fibre surface is taken as 50 Å thick, the heats just calculated, if reckoned per gram of surface material, are 2.8 x 10 -3 cal and — 2.4 x 10 -3 cal, respectively. The former is about the same as the heat produced per gram in a muscle twitch. During the passage of an impulse there is known to be an interchange of Na and K ions between the axoplasm and the outside fluid. When isotonic solutions of NaCl and KCl are mixed there is a production of heat. A substantial part of the heat during an impulse may be derived from the interchange of Na and K. Another part may be associated with chemical reactions occurring in the excitable membrane during the cycle of permeability change accompanying the passage of an impulse. The negative heat production is discussed. It cannot be connected with ‘pumping back’ the Na and K ions; this is a much slower process and anyhow would probably involve a positive heat production. It may be a sign of endothermic chemical reactions, representing a first (anaerobic) stage in recovery, which occur in the surface membrane following the completion of the permeability cycle. The question is considered whether the positive and negative phases of the heat production could be due to the discharge and recharge, during the action potential, of the condenser residing in the excitable membrane. The heats so calculated are of the right order of size, but on present evidence the time relations seem to be quite wrong. The amount of K which escapes per impulse from Maia nerve during slow repetitive stimulation at 0° C was measured. It depends greatly on frequency of stimulation; at ‘zero frequency’ it was about 9 X 10 -8 mole/g x impulse.

The priority of the heat production in a muscle twitch

1958 ◽  
Vol 148 (932) ◽  
pp. 397-402 ◽  
Keyword(s):  
Chemical Reactions ◽  
Heat Production ◽  
Mechanical Response ◽  
Substantial Part ◽  
Hypertonic Solution ◽  
Isotonic Solution ◽  
Muscle Twitch ◽  
Previous Conclusion

When a muscle has been soaked in a moderately hypertonic solution its mechanical response to a shock is delayed, but its heat production is almost normal and starts considerably earlier than its shortening. After a more hypertonic solution the mechanical response is abolished, but a substantial part of the heat production remains. These effects are rapidly reversed by soaking in a normal isotonic solution. They strengthen the previous conclusion that chemical reactions triggered by a stimulus precede the mechanical response.

Linear electrical properties of striated muscle fibres observed with intracellular electrodes

1964 ◽  
Vol 160 (978) ◽  
pp. 69-123 ◽  
Keyword(s):  
Electrical Properties ◽  
Striated Muscle ◽  
Surface Membrane ◽  
Fibre Surface ◽  
Step Function ◽  
Muscle Fibres ◽  
Appreciable Effect ◽  
Current Application ◽  
Current Path ◽  
Wide Range

The linear electrical properties of muscle fibres have been examined using intracellular electrodes for a. c. measurements and analyzing observations on the basis of cable theory. The measurements have covered the frequency range 1 c/s to 10 kc/s. Comparison of the theory for the circular cylindrical fibre with that for the ideal, one-dimensional cable indicates that, under the conditions of the experiments, no serious error would be introduced in the analysis by the geometrical idealization. The impedance locus for frog sartorius and crayfish limb muscle fibres deviates over a wide range of frequencies from that expected for a simple model in which the current path between the inside and the outside of the fibre consists only of a resistance and a capacitance in parallel. A good fit of the experimental results on frog fibres is obtained if the inside-outside admittance is considered to contain, in addition to the parallel elements R m = 3100 Ωcm 2 and C m = 2.6 μF/cm 2 , another path composed of a resistance R e = 330 Ωcm 2 in series with a capacitance C e = 4.1 μF/cm 2 , all referred to unit area of fibre surface. The impedance behaviour of crayfish fibres can be described by a similar model, the corresponding values being R m = 680 Ωcm 2 , C m = 3.9 μF/cm 2 , R e = 35 Ωcm 2 , C e = 17 μF/cm 2 . The response of frog fibres to a step-function current (with the points of voltage recording and current application close together) has been analyzed in terms of the above two-time constant model, and it is shown that neglecting the series resistance would have an appreciable effect on the agreement between theory and experiment only at times less than the halftime of rise of the response. The elements R m and C m are presumed to represent properties of the surface membrane of the fibre. R e and C e are thought to arise not at the surface, but to be indicative of a separate current path from the myoplasm through an intracellular system of channels to the exterior. In the case of crayfish fibres, it is possible that R e (when referred to unit volume) would be a measure of the resistivity of the interior of the channels, and C e the capacitance across the walls of the channels. In the case of frog fibres, it is suggested that the elements R e , C e arise from the properties of adjacent membranes of the triads in the sarcoplasmic reticulum . The possibility is considered that the potential difference across the capacitance C e may control the initiation of contraction.

Address of the President Sir Alan Hodgkin, O. M. at the Anniversary Meeting, 30 November 1973

1974 ◽  
Vol 185 (1078) ◽  
Keyword(s):  
Surface Membrane ◽  
Muscular Activity ◽  
Nerve Impulse ◽  
Muscular Contraction ◽  
Single Fibre ◽  
Muscle Fibres ◽  
Constant Length ◽  
Axon Membrane ◽  
Ultrastructural Studies ◽  
Mathematical Skill

The Copley Medal is awarded to Professor A. F. Huxley, F. R. S. A. F. Huxley has made outstanding contributions to our knowledge of the nerve impulse and of the mechanism by which muscle fibres are caused to contract. Jointly with Hodgkin, he introduced the powerful method of intracellular recording from nerve cells and showed that during the propagation of an impulse the mem­brane potential reverses its sign, and does not simply fall towards zero as had been widely believed. This work - interrupted by the 1939-45 war, but later resumed - led to the proposal that the impulse arises from a transient influx of sodium ions through the axon membrane. The ‘ionic theory’ of nervous conduction was then established by a series of convincing experiments and calculations for which Huxley later shared the Nobel Prize. Huxley next turned his attention to the mechanism of muscular contraction. He equipped himself for this purpose by inventing a new type of interference microscope. In experiments on living isolated muscle fibres, Huxley showed that contraction is accompanied by a shortening of the isotropic band of each sarco­mere, while the remaining portion (the anisotropic band) retains approximately constant length. His findings complemented the important ultrastructural studies of H. E. Huxley and led them both to propose a ‘sliding filament’ mechanism as the basis of muscular motion. During further microscopic observations on the living muscle fibre, A. F. Huxley produced most striking evidence on the way in which an excitatory potential change of the surface membrane is communicated, through local tubular channels, to the interior of the fibre where it activates the contractile elements. In his most recent work, A. F. Huxley has continued to develop his single-fibre technique to resolve even finer details of the dynamic changes which occur during muscular activity. His work is characterized by a rare combination of profound theoretical insight, mathematical skill and superb technical mastery, all of which has enabled him to select problems of first-rate importance and to pursue them with outstanding success.

Body Temperature and Singing in the Bladder Cicada, Cystosoma Saundersii

1979 ◽  
Vol 80 (1) ◽  
pp. 69-81 ◽  
Author(s):  
R. K. JOSEPHSON ◽  
D. YOUNG
Keyword(s):  
Heat Production ◽  
Cell Biology ◽  
Sound Production ◽  
Marked Decrease ◽  
Cooling Curve ◽  
Muscle Twitch ◽  
Temperature Excess ◽  
Air Mixing ◽  
Artificial Heat ◽  
Sound Pulses

1. Body temperatures during singing were measured in the cicada, Cystosoma saundersii Westwood, both in the field and in tethered animals indoors. 2. The temperature of the sound-producing tymbal muscle rises rapidly during singing to reach a plateau approximately 12°C above ambient. This produces a temperature gradient in the abdominal air sac which surrounds the muscle. When singing stops, the tymbal muscle cools exponentially. 3. Heat production during singing, estimated from the cooling curve, is 4.82 cal min−1 g muscle−1. Generation of the same temperature excess in the air sac by an artificial heat source yields an estimated heat production of 54.4 cal min−1 g muscle−1. This discrepancy may be caused by air mixing in the air sac during singing. 4. As temperature rises, tymbal muscle twitch contractions become faster and stronger. This and heat transfer to the thorax cause changes in the song pattern: a marked decrease in the interval between the two sound pulses produced by a single tymbal buckling and a lesser decrease in the interval between bucklings. The fundamental sound period remains unaltered. These effects are consistent with earlier data on sound production. Note: Present address: Department of Developmental and Cell Biology, University of California, Irvine, California 92717, U.S.A.

Heat production in oscillating chemical reactions: three examples

1987 ◽  
Vol 112 (1) ◽  
pp. 95-100 ◽  
Author(s):  
I. Lamprecht ◽  
B. Schaarschmidt ◽  
T. Plesser
Keyword(s):  
Chemical Reactions ◽  
Heat Production

Richard Hume Adrian, D. L. 2nd Baron Adrian of Cambridge. 16 October 1927—4 April 1995

1997 ◽  
Vol 43 ◽  
pp. 15-30
Author(s):  
Andrew Huxley
Keyword(s):  
Education Reform ◽  
Major Part ◽  
Striated Muscle ◽  
Surface Membrane ◽  
Scientific Work ◽  
Fibre Surface ◽  
Electrical Behaviour ◽  
Cambridge University ◽  
Vice Chancellor ◽  
House Of Lords

Richard Adrian was a distinguished electrophysiologist who cleared up many of the puzzles about the electrical behaviour of striated muscle. The situation in this tissue is exceptionally complex, both because the surface membrane is invaginated to form the transverse tubules, whose surface area is several times that of the fibre surface, and because substantial amounts of charge are carried across these membranes by processes connected with the turning on of contraction itself. Apart from his scientific work, he played an important part in the life of Cambridge University, as Master of Pembroke College for 11 years and as Vice-Chancellor for two. In the House of Lords, he contributed in an important way to the debates leading up to the new Act of 1986 governing experiments on living animals, and took a major part in the defence of academic freedom in the universities when it was threatened in the Education Reform Bill of 1988.

Primary and Secondary Regulation of Quantal Transmitter Release: Calcium and Sodium

1980 ◽  
Vol 89 (1) ◽  
pp. 5-18
Author(s):  
RAMI RAHAMIMOFF ◽  
AHARON LEV-TOV ◽  
HALINA MEIRI
Keyword(s):  
Action Potential ◽  
Nerve Terminal ◽  
Transmitter Release ◽  
Surface Membrane ◽  
Nerve Impulse ◽  
High Frequency Stimulation ◽  
Extracellular Medium ◽  
Presynaptic Nerve Terminal ◽  
Virtual Absence ◽  
Frequency Stimulation

Calcium is the prime regulator of quantal acetylcholine liberation at the neuromuscular junction; its entry through the presynaptic membrane and the level of free [Ca]ln most probably determine the number of transmitter quanta liberated by the nerve impulse. The level of free [Ca]ln, in turn, is controlled by a number of subcellular elements: mitochondria, endoplasmic reticulum, vesicles, macromolecules and the surface membrane. The action potential induced calcium entry is not the only factor responsible for coupling nerve terminal depolarization with increased transmitter release; increased transmitter release occurs also in the virtual absence of calcium ions in the extracellular medium, when a reversed electrochemical gradient for calcium probably exists during action potential activity. Several lines of evidence suggest that the entry of sodium ions is responsible for this augmented transmitter release: the tetanic potentiation observed under reversed calcium gradient is blocked by tetrodotoxin; tetanic and post-tetanic potentiation are augmented and prolonged by ouabain; the amplitude of the extracellular nerve action potential is reduced with high-frequency stimulation, in parallel with increased spontaneous quantal release. In addition, sodium-filled egg-lecithine liposomes augment quantal liberation. The augmentory effect of sodium on transmitter release is probably due to an intracellular calcium translocation, since no preferred timing after the action potential is observed. Thus the level of [Na]ln in the presynaptic nerve terminal can control indirectly the efficiency of synaptic transmission.

A note on the heat of activation in a muscle twitch

1950 ◽  
Vol 137 (888) ◽  
pp. 330-331 ◽  
Keyword(s):  
Heat Production ◽  
Mechanical Work ◽  
Muscle Twitch ◽  
Extreme Loads

Under extreme loads a stimulated muscle neither shortens nor develops tension. The heat production in a twitch is then rather less than one-half of its value with maximal shortening. Under such conditions the heat of shortening and the mechanical work are nil. The remaining heat, therefore, is heat of activation alone. It is about the same in magnitude and onset as the heat of activation at ordinary lengths.

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