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Original Articles

Bernstein's Long Path to Membrane Theory: Radical Change and Conservation in Nineteenth-Century German Electrophysiology

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Pages 306-337 | Published online: 17 Oct 2011
 

Abstract

This article aims at illustrating the historical circumstances that led Julius CitationBernstein in 1902 to formulate a membrane theory on resting current in muscle and nerve fibers. It was a truly paradigm shift in research into bioelectrical phenomena, if qualified by the observation that, besides Bernstein, many other electrophysiologists between 1890 and 1902 borrowed ideas from the recent ionistic approach in the physical-chemistry domain. But Bernstein's subjective perception of that paradigm shift was that it constituted a mere reinterpretation of the so-called preexistence theory advanced by his teacher Emil du Bois-Reymond in the first half of the nineteenth century.

Acknowledgments

We wish to express our deep gratitude to Dr. Stanley Finger for his suggestions and for his revision of our manuscript and to Dr. Marco Piccolino for his help with the illustrations. The work on which this article is based was partially funded by an Italian National Grant (PRIN 2007).

Notes

1From this point of view Timothy Lenoir proves to be lacking in the historical background information in his essay (1986).

2In 1907, Boruttau was appointed professor of physiology at the University of Berlin.

3For the first exposition of the Alterationstheorie, see CitationHermann (1867). However, the term Demarcationsstrom was introduced only later (for example 1879, p. 236).

Figure 5. Ludimar Hermann (1838–1914), a student of du Bois-Reymond who strongly criticized his teacher's contention of the existence of a measurable current at the surface of intact nerves or muscles at rest. In the polemics against Hermann, Bernstein joined du Bois-Reymond's camp.

Figure 5. Ludimar Hermann (1838–1914), a student of du Bois-Reymond who strongly criticized his teacher's contention of the existence of a measurable current at the surface of intact nerves or muscles at rest. In the polemics against Hermann, Bernstein joined du Bois-Reymond's camp.

4About the dispute involving du Bois and his pupils Bernstein and Hermann, see CitationFinkelstein (2006).

5 CitationBernstein (1912, p. 5, footnote).

7This essay was reprinted as the first chapter of CitationBernstein (1871).

Figure 7. The title page of the 1871 extensive monograph in which Bernstein reported the results of his electrophysiological investigation with the differential rheotome.

Figure 7. The title page of the 1871 extensive monograph in which Bernstein reported the results of his electrophysiological investigation with the differential rheotome.

8Replying to the criticism raised against du Bois-Reymond's molecular theory, Bernstein explained that “in the hypothesized form, the molecules are not intended as being the real ones but rather only representative of forces which we imagine under their appearance” (1874, p. 58).

9Mistakenly CitationLenoir (1986, p. 34) dated this theory to 1871.

10See also CitationNernst (1889). For a reconstruction of Nernst's work and his scientific environment, see CitationBarkan (1999).

11A biographical sketch framing Traube as a biochemist is offered in CitationSourkes (1955).

12For reasons that will be clear later in the article, we have reported, with some minor changes, the equation presented by Nernst's student Max CitationLe Blanc (1896, p. 156).

13This concept was anticipated in CitationNernst (1897, p. 615, Note 3), where it is stated that the physiological effect of alternating current is caused by “variations in concentration at the cellular membranes (impermeable against certain electrolytes, i.e., ions).”

14W. J. Tschgowetz in the German transliteration.

15Research conducted over a five-year period starting in 1890 on living animal and plant cells. See CitationDe Weer (2000).

16But Des Coudres' name appears only in CitationBoruttau (1904, p. 433).

17Strong intended to establish the conducting function of nerves based only on their cable-like structure, without attributing any function to the membrane, except as an isolating sheath.

18In his 1902 article, Bernstein mentioned monobasic potassium phosphate (KH2PO4) as an electrolyte but later (1912, p. 94) mentioned bibasic potassium phosphate as the inorganic salt present in the muscle in which potassium binds to phosphoric acid.

19For the experiments with tap water Macdonald used to cause the extreme case of injury, see the expanded version (1902). For the concentration cells Macdonald referred to see CitationLe Blanc's manual (1896) and the English edition of CitationNernst's treatise (1923).

20The term “preexistence” was coined by du Bois-Reymond in 1867 during the controversy with Hermann. See du Bois-Reymond (1867/ 1877, p. 359): “die Präexistenz des elektrischen Gegensatzes in den Muskeln und Nerven.” See also CitationMunk (1868).

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