Bioelectricity  ·  Article

Ion exchange as the cause of contraction and rhythmic activity

In The Dynamics of Living Matter (1906), Jacques Loeb proposed that contraction in muscle or nerve is caused by an exchange of sodium or potassium for calcium or magnesium in certain compounds.1 He reached the idea from the timing of rhythmical contractions in salt solutions.2 He expected the exchange to change a physical property of the compound, such as surface tension.3

Earliest held
1888, Binet, A.
Most discussed in
Researches on Irritability of Plants, 1913
In the library
332 passages in 22 works
Rewritten
2026-11-05
01

Loeb's proposal

Loeb noted that a slight amount of acid added to a NaCl solution shortens the latent period before rhythmical contractions begin.2 He suggested that the acid may act like the salts he had discussed, by liberating calcium from certain organic combinations in the muscle or nerve.2 Because CO2 is produced in the muscle itself, he judged this relation to be of importance.2 From these facts he drew the idea that contraction follows an exchange of Na or K for Ca or Mg, or the reverse.1 Loeb offered the idea as a suggestion drawn from these facts, not as a measured result.1

02

Physical change in the compound

Loeb argued that the exchange must be accompanied by a change in some physical property of the compound. His examples were surface tension, state of aggregation, viscosity and absorbing power for water.3 He held that such a change may determine or facilitate contraction.3 Loeb named these properties only as possible examples and did not identify which one changes. The latent period served as the observable link between the salt solutions and the idea.2

03

Latent period in other work

Other physiologists, such as Bose and Bayliss, also measured latent periods. Bose recorded latent periods in the plant Mimosa with a Resonant Recorder able to resolve a thousandth of a second.7 In 1913 he reported that the period is shorter under stronger stimulus but constant above a maximal one.5 In 1927 he reported that fatigue lengthened it from 0.10 to 0.14 second.8 A 1907 record showed a latent period falling from 25 to 20 minutes on repeated stimulus.4 Bayliss gave a latent period of 0.023 second for the cephalopod eye responding to light.6

04

Cautions on the term

Pavlov raised a warning about reading the latent period too simply. He wrote that what his own experiments had often called the latent period of a conditioned reflex was in fact a delay.9 He called it a preliminary inhibitory period and said it was certainly not the true latent period of the reflex.9 Pavlov's caution was not directed at Loeb.

SourcesEach quotation was checked word for word against the passage it opens.
  1. These facts suggested the idea that the process of contraction is caused by an exchange of Na or K for Ca or Mg, or vice versa, in cer- tain compounds in the muscle (or nerve).Loeb, J., 1906 · The Dynamics of Living Matter · open at passage 222
  2. The addition of a slight amount of acid to a - NaCl solution shortens the latent period for the beginning of the rhythmical contractions.Loeb, J., 1906 · The Dynamics of Living Matter · open at passage 222
  3. a change in some physical property of the compound, e.g. the surface tension, the state of aggregation, viscosity, absorbing power for water, etc.Loeb, J., 1906 · The Dynamics of Living Matter · open at passage 222
  4. Lower record shows latent period to be 25 minutes. After re-balance, stimulus of 5 seconds was again applied. Upper record now shows reduction of latent period to 20 minutesBose, J. C., 1907 · Comparative Electro-Physiology: A Physico-Physiological Study · open at passage 943
  5. The latent period is in general shorter under stronger intensity of stimulus; but the value becomes constant above a maximal stimulus.Bose, J. C., 1913 · Researches on Irritability of Plants · open at passage 746
  6. Eye of Cephalopod (Eledone). Simple positive variation with light, after latent period of 0'023 secondBayliss, W. M., 1915 · Principles of General Physiology · open at passage 2247
  7. The latent period of the pulvinus may be determined with great accuracy by means of the Resonant Recorder, by which it is possible to estimate time-intervals as short as a thousandth part of a second.Bose, J. C., 1926 · The Nervous Mechanism of Plants · open at passage 171
  8. The second record was taken before the plant had recovered from its previous shock and was, therefore, in a state of fatigue. The latent period was now prolonged from o.io to 0.14 second.Bose, J. C., 1927 · Plant Autographs and Their Revelations · open at passage 52
  9. which could appro- priately be termed the “ preliminary inhibitory period,” but which is certainly not the true latent period of the reflex.Pavlov, I. P., 1927 · Conditioned Reflexes: An Investigation of the Physiological Activity of the… · open at passage 262
Linked ideas
Loeb's salt solutions, where Na, K, Ca and Mg act against one another, are the setting for the exchange idea.
Both tie muscle behaviour to the state of compounds in the tissue and their physical properties such as water uptake.
Bose's measurements of latent period under fatigue and stimulus strength bear on how excitable tissue responds.
Bose found latent period shortens with stimulus strength up to a maximal stimulus, a graded relation of the same kind.
Conditioned reflexin tension with
Pavlov cautioned that an apparent latent period in conditioned reflexes was a delay, which complicates its use as evidence.
Later ion-based accounts of excitation concern the same Na, K and Ca balance, though the held Loeb passage does not speak of membranes.
Where it is discussedPassages matching exchange of Na or K for Ca or Mg, rhythmical contractions in NaCl, liberating calcium from organic combinations, latent period
1913Researches on Irritability of Plants · Bose, J. C.95
1906Plant Response as a Means of Physiological Investigation · Bose, J. C.43
1927Conditioned Reflexes: An Investigation of the Physiological Activity of the… · Pavlov, I. P.33
1926The Nervous Mechanism of Plants · Bose, J. C.30
1918Life Movements in Plants · Bose, J. C.24
1907Comparative Electro-Physiology: A Physico-Physiological Study · Bose, J. C.21
1928The Motor Mechanism of Plants · Bose, J. C.16
1915Principles of General Physiology · Bayliss, W. M.14
1906The Integrative Action of the Nervous System · Sherrington, C. S.12
1927Plant Autographs and Their Revelations · Bose, J. C.8