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Ion Gradients and Movements in Excitable Tissues

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Moving Questions

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Abstract

When certain tissues, such as nerve and muscle, are perturbed in specific ways they change their properties in a characteristic manner. They are “excitable tissues,” and when stimulated they respond. A nerve trunk stimulated electrically may cause the innervated muscle to shorten: something has traveled from the point of stimulation to and then along the muscle, inducing contraction.

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Notes to Chapter 5

  1. For relevant historical accounts see Brazier (1959); Hille (1992); Hodgkin (1992); and Tasaki (1959).

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  2. Translations quoted are from Weidmann (1955), p. 116, and Glynn (1989a), p. 39.

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  3. Young (1936).

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  4. Curtis and Cole (1938). They acknowledged Young’s introducing them to squid giant axons.

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  5. Cole and Curtis (1939). R. L. Post (personal communication, 1995) pointed out that Cole and Curtis could have derived the membrane currents from their measurements, applying their analysis of cable properties, by taking derivatives of the action potentials. He quoted Curtis as not having had sufficient confidence in those measurements to make such analyses then.

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  6. Cole and Hodgkin (1939).

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  7. The resting membrane potential is expressed relative to the potential of the bathing medium defined as 0 mV; it has then a negative sign. The action potential magnitude is expressed as an absolute number, relative to the membrane potential defined now as 0 mV. Algebraic addition of the resting potential and the action potential magnitude gives the overshoot magnitude.

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  8. Hodgkin and Huxley (1939). They too thanked Young for his contribution.

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  9. Curtis and Cole (1942).

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  10. Ibid., pp. 136, 141, 142.

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  11. Hodgkin (1992), p. 252.

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  12. This calculation is based on intracellular concentrations subsequently reported by Steinbach and Spiegelman (1943); with values available previously (Bear and Schmitt, 1939) the maximal value would be +14 mV.

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  13. Hodgkin and Huxley (1945).

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  14. Höber (1946), p. 388.

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  15. Grundfest (1947).

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  16. Hodgkin and Huxley (1947).

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  17. Ibid., pp. 341–342.

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  18. Katz (1947).

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  19. Hodgkin and Katz (1949), pp. 73–74; Hodgkin et al. (1949).

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  20. Hodgkin et al. (1949).

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  21. Hodgkin and Katz (1949), p. 65.

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  22. Hodgkin et al. (1949).

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  23. Goldman (1943).

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  24. Hodgkin and Katz (1949).

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  25. Hodgkin et al. (1949).

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  26. Cole (1949); Marmont (1949).

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  27. Hodgkin et al. (1952); Hodgkin and Huxley (1952a,b,c,d).

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  28. Stationary action potentials were obtained with electrodes clamping potentials over appreciable lengths of axons (a “space clamp”).

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  29. Cole (1954) and Cole et al. (1955) reported the failure to replicate Hodgkin and Huxley’s solution; Cole (1958) retracted those reports.

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  30. Hodgkin and Huxley (1952d), p. 541.

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  31. Ibid., pp. 543–544.

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  32. Webb and Young (1940); Hodgkin and Huxley (1939); Curtis and Cole (1942).

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  33. Grundfest (1950).

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  34. Cole (1968), p. 145.

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  35. Lorente de N6 (1944).

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  36. Lorente de N6 (1947).

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  37. Feng and Gerard (1930). The quotation is from Feng and Liu (1949), p. 1.

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  38. Lorente de N6 (1947), as quoted in Feng and Liu (1949), p. 1.

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  39. Feng and Liu (1949).

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  40. Grundfest (1950), pp. 17–22.

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  41. For example, Lorente de N6 (1947) proposed that “K+ and Na + can play a specific role only in so far as they directly or indirectly participate in enzymatic reactions“ (p. 107), with the membrane potential then ”referable to the oxidation-reduction systems of the chain of oxidative enzymes“ (p. 106).

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  42. Keynes (1948, 1949a). Keynes used cuttlefish axons because cuttlefish, unlike squid, could be kept in Cambridge.

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  43. Keynes and Lewis (1951a); Keynes (1951a). The incremental flux, due to the action potential, was calculated to be greater than 2.3 x 10-’ coulombs per impulse, whereas repolarizing the membrane required only 1.4 x 10-7 coulombs.

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  44. Keynes (1951b); Keynes and Lewis (1951b).

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  45. Keynes and Lewis (1951b), p. 179.

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  46. Rothenberg and Feld (1948); Rothenberg (1950).

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  47. Keynes and Lewis (1951a).

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  48. Boyle and Conway (1941).

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  49. Hodgkin and Huxley (1946), p. 377.

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  50. Hodgkin and Huxley (1947), p. 364.

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  51. Hodgkin (1951).

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  52. Hodgkin and Huxley (1947), p. 365.

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  53. Hodgkin and Huxley (1952d), p. 541.

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  54. Keynes (1951a,b).

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  55. Huxley and Stämpfli (1949), p. 339.

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  56. Huxley and Stämpfli (1951a,b).

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  57. Graham and Gerard (1946).

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  58. Ling and Gerard (1949a).

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  59. Nastuk and Hodgkin (1950). They thanked Ling for “demonstrating his experimental technique” (p. 72).

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  60. Woodbury et al. (1950).

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  61. Draper and Weidman (1951).

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  62. Biographical information is from interviews, reminiscences (Hodgkin, 1992; Keynes, 1989), and biographies (Huxley, 1992).

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  63. Hodgkin (1976), p. 11.

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  64. Solomon (1989), p. 129.

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  65. Hodgkin (1992).

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  66. Hodgkin (1949); Hodgkin et al. (1949); Cole (1949). Cole’s paper is also notable for its contrasting focus on electronic currents to the neglect of ionic currents.

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  67. Expressing nerve activity in terms of equivalent circuits was not a novel idea (see, for example, Rushton, 1937).

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  68. Some ideas about these structures were, however, often far from today’s proposed structures; see Cole (1968), pp. 537–539, for conceptions of that period.

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  69. Hodgkin and Huxley (1945), p. 192.

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  70. See, for example, Robinson (1986a, 1992).

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© 1997 American Physiological Society

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Robinson, J.D. (1997). Ion Gradients and Movements in Excitable Tissues. In: Moving Questions. People and Ideas Series. Springer, New York, NY. https://doi.org/10.1007/978-1-4614-7600-9_5

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  • DOI: https://doi.org/10.1007/978-1-4614-7600-9_5

  • Publisher Name: Springer, New York, NY

  • Online ISBN: 978-1-4614-7600-9

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