Summary of the major innovations of Emil du Bois-Reymond (1818-1896

Finkelstein, Gabriel

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Mechanical Neuroscience: Emil du Bois-Reymond’s Innovations in Theory and Practice

Gabriel Finkelstein

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Mechanical neuroscience: Emil du Bois-Reymond's innovations in theory and practice

Summary of the achievements of Emil du Bois-Reymond (1818-1896

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OPINIONpublished: 30 September 2015doi: 10.3389/fnsys.2015.00133Frontiers in Systems Neuroscience | www.frontiersin.org 1 September 2015 | Volume 9 | Article 133Edited by:Mikhail Lebedev,Duke University, USAReviewed by:Marina Bentivoglio,University of Verona, Italy*Correspondence:Gabriel Finkelsteingabriel.finkelstein@ucdenver.eduReceived: 08 September 2015Accepted: 14 September 2015Published: 30 September 2015Citation:Finkelstein G (2015) Mechanicalneuroscience: Emil duBois-Reymond’s innovations in theoryand practice.Front. Syst. Neurosci. 9:133.doi: 10.3389/fnsys.2015.00133Mechanical neuroscience: Emil duBois-Reymond’s innovations intheory and practiceGabriel Finkelstein *Department of History, University of Colorado Denver, Denver, CO, USAKeywords: Emil du Bois-Reymond, neuroscience, paradigm, history, nineteenth-centuryIntroductionContemporary neuroscience unites three scientific traditions: one in anatomy that goes backto the Greeks (functional localization), one in chemistry that arose in the twentieth-century(neurotransmitters), and one in physics that arose in the nineteenth-century (electrical signals)(Clarke and Jacyna, 1987; Finger, 1994; Valenstein, 2005; McComas, 2011). Emil du Bois-Reymondis the father of the last. He’s not quite Victor Frankenstein, but he’s close.When du Bois-Reymond began his investigations neurophysiologists were divided. Somefollowed the physician Luigi Galvani (1737–1798) in ascribing the action of nerves and muscles tovital powers of “animal electricity.” Others sided with the physicist Alessandro Volta (1745–1827)in believing that the muscular contractions that Galvani observed were an artifact of electricitygenerated by the contact of metal with organic tissue (Pera, 1992; Piccolino, 1998; Piccolino andBresadola, 2013). Recognizing value in both positions, du Bois-Reymond solved the problem ofcontact electricity, set forth a program of biological reduction, and demonstrated the electricalnature of nerve signals. In a little less than two years—from March, 1841 to January, 1843—hecreated the discipline of electrophysiology.That’s the short version of du Bois-Reymond’s innovation. We know that it’s true becausewinners write history—at least, that’s what we’ve been told. In the case of du Bois-Reymond,however, the truth is closer to this: those who write history win. Does this fact change his story?Consider the evidence.TheoryEmil du Bois-Reymond was born in Berlin on 7 November 1818 to family of Huguenot origins(Finkelstein, 2013). His mother’s forebears were merchants, scholars, and artists that had livedin Berlin for generations; his father was a poor, self-taught immigrant from the Swiss canton ofNeuchâtel. Du Bois-Reymond grew up speaking French, reading the philosophes, and chafing atthe confines of his father’s Calvinism. His first impulse on entering university was to embraceRomanticism, but it didn’t take long for him to ditch Naturphilosophie in favor of a mechanicalview of the world. Clues to his conversion can be found in his reading: Lucretius in the summer of1838, French physics shortly thereafter, and the Cours de philosophie positive of Auguste Comte—most likely—in 1842. The result, as he reported to a friend, regarded science from a perspective thatwas nearly Cartesian:I have sworn to uphold the truth that no forces operate in the organism other than those commonto physics and chemistry; that, where these do not suffice in explanation by means of the mathematico-physical method, one must either look at a specific case of the force in question, or one must assumenew forces, which, of the same order as the physico-chemical inherent in matter, always reduce to onlyattractive and repulsive components (du Bois-Reymond, 1918, p. 108).Finkelstein Mechanical neuroscienceIt should be noted that du Bois-Reymond’s materialism waspurely methodological. As he saw it, force and matter servedas terms of convenience, and anyone who confused themwith real entities was committing the same error as primitiveancestors who “once peopled bush and fountain, rock, air, andsea with creatures of their imagination” (du Bois-Reymond,1848–1884, pp. xl–xli). Anthropomorphic fantasies had no placein science.With respect to biology du Bois-Reymond’s outlook translatedinto a repudiation of vitalism. To be sure, he was not thefirst German to adopt this position—the Romantic psychiatristJohann Christian Reil (1759–1813) had done so in 1796, ashad the psychologist Hermann Lotze (1817–1881) in 1842—but du Bois-Reymond was the first to effect a lasting changein the outlook of his colleagues. It was one thing fortheoreticians like Reil and Lotze to espouse a particularview of nature, and it was altogether another for a scientistlike du Bois-Reymond. This is why his words carried somuch weight. Looking back at the development of hisdiscipline over the course of the nineteenth-century, theBelgian physiologist Paul Heger (1846–1925) deemed duBois-Reymond’s pronouncement “an act of courage whoseworth is difficult to appreciate today” (Heger, 1896–1897, p.565).In this regard the three elements of du Bois-Reymond’spositivism (biological reduction, functional analysis, andmethodological materialism) constitute a shift in the paradigmof neuroscience. Very few neuroscientists still endorse the crudematerialism that Carl Vogt (1817–1895), Jacob Moleschott(1822–1893), and Ludwig Büchner (1824–1899) popularized intheir polemics against religion in the 1850s (Gregory, 1977).By contrast, most share du Bois-Reymond’s opinion that vitalpowers do not operate in biology, that explanations of thenervous system require accounts of the chemical and physicalprocesses that drive it, and that certain problems, like theultimate essence of matter, consciousness, and free will, liebeyond the understanding of science.PracticePerhaps the greatest of du Bois-Reymond’s innovations wasan experimental design that solved three problems that vexedthe study of animal electricity. First, du Bois-Reymond devisedneutral means of coupling instruments to tissue, most notably“non-polarizable” electrodes formed from an amalgam ofzinc, zinc sulfate, and modeling clay. Second, he inventeddevices like the “magneo-electrometer” (AC generator) andthe “rheocord” (potentiometer) that delivered graded shocksto his preparations. Last, he constructed a galvanometersensitive enough to record the results of his protocols. Thesebreakthroughs allowed him to detect action currents in frogmuscles in 1843; 4 years later the addition of a Wheatstonebridge circuit to his set-up let him to demonstrate the sameelectrical signals in human subjects (see Figure 1) (Finkelstein,2013).Du Bois-Reymond’s most famous experiment proceeded asfollows. He attached the leads of his galvanometer to platinumFIGURE 1 | Du Bois-Reymond’s demonstration of voluntary tetaniccurrent. The arrows indicate the flow of current through the body. Thegalvanometer detecting the signal rests on a separate shelf to avoiddisturbance. Reprinted from du Bois-Reymond (1848–1884), Volume 2, part 2,plate V, Figure 147.plates resting in “conducting vessels” filled with saline. Afterimmersing his fingers in the electrolyte, he waited for thegalvanometer needle to come to rest. All of a sudden, he tensedone of his arms. The needle jumped. Observers in Berlin, Paris,and London were astonished by his exhibition; indeed theperformance, which was simple and striking, made his name asmuch as any of his scholarly publications. The point was notlost on du Bois-Reymond, who went on to devise a panoplyof demonstration aids like “the twitch telegraph,” “the frogalarm,” “the frog pistol,” “the mirror multiplier,” and “the feathermyograph,” many of which still feature in the collections ofscience museums today.The significance of du Bois-Reymond’s instruments escapedhis French and English colleagues at first, but within a generationeven du Bois-Reymond’s skeptics adopted his methods. In fact,it wouldn’t be that much of a stretch to describe the historyof neuroscience in the image of du Bois-Reymond: a march oftechnological progress from the galvanometer to the optogeneticsensor, all originating in table-top experiments that he carried outwith apparatus he built himself.Du Bois-Reymond’s instrumental methods embodied apositivist commitment to measurement that permanentlychanged the practice of science (Kuhn, 1961, p. 190). LordKelvin expressed his attitude perfectly: “I often say that whenyou can measure what you are speaking about, and expressit in numbers, you know something about it; but when youcannot measure it, when you cannot express it in numbers,your knowledge is of a meager and unsatisfactory kind: itmay be the beginning of knowledge, but you have scarcely inFrontiers in Systems Neuroscience | www.frontiersin.org 2 September 2015 | Volume 9 | Article 133Finkelstein Mechanical neuroscienceyour thoughts advanced to the stage of science” (Thompson,1889, p. 73). It might be argued that our knowledge of thebrain remains meager and unsatisfactory, that there is stillno consensus about what to measure, at what scale, andover what span of time. All the same, Alan Lloyd Hodgkinand Andrew Huxley’s mathematical model of neuron actionpotentials endures as one of the great achievements of twentieth-century physiology. Had du Bois-Reymond been alive to hearthe news of their Nobel Prize, he would have endorsed itimplicitly.CommunicationNone of du Bois-Reymond’s innovations would have had quitethe same effect if he hadn’t advertised them to colleagues,students, and the public. Fortunately, he possessed an exceptionaltalent at communication. He helped found the Berlin Society ofPhysics and the Berlin Society of Anthropology, organizationsthat served as sounding boards for his discoveries; he workedas permanent secretary of the Prussian Academy of Sciences,rector of the University of Berlin, and dean of the medicalschool, positions that required him to deliver formal addresses;and he edited the leading German journal of physiology, apost that helped him shape the discipline. He also directedPrussia’s first institute of physiology, instructed medical studentsin animal electricity, and taught Berlin’s most popular class.Finally, he conducted demonstrations in Berlin, Paris, andLondon, spoke to audiences throughout Germany and theNetherlands, and reprinted his speeches in the DeutscheRundschau, the Revue scientifique, Nature, and the PopularScience Monthly.Like all great performers du Bois-Reymond thought hardabout his craft. He compared professors who spoke poorlyto eccentrics who wandered about in their dressing-gowns(du Bois-Reymond, 1912, p. 492). Remarking on one publicsession at the Academy of Sciences, he told his wife, “Theonly decent address was mine, of course. . . profound, allusive,brief, masterfully arranged. Caviar for the audience” (du Bois-Reymond, 1882). He used the same idiom to describe thelectures of his tour of the Rhineland: “Apparently no oneis aware that you can present a scientific discourse with agarnish of poetic reflections and numerical figures, like afilet with mixed pickles and olives. In Cologne they insistedthat I had read from a manuscript” (du Bois-Reymond,1877).Du Bois-Reymond’s attention to rhetoric was deliberate. Earlyin his career he realized that “popularizers of science persist in thepublic mind as memorial stones of human progress long after thewaves of oblivion have surged over the originators of the soundestresearch” (du Bois-Reymond, 1912, p. 354). He never abandonedhis investigations of animal electricity, but he alsomade certain toaddress his audience. “The time of poetic production in Europeannations appears to have passed,” he wrote to his fiancée in 1853,“and talent, which might otherwise have achieved somethingthere, throws itself into oratory and journalism, and later dabblesin politics” (du Bois-Reymond, 1853). He was somewhat blunterabout the change two decades later. “Today Goethe would beholding forth at the Reichstag” (du Bois-Reymond, 1912, p.607).Du Bois-Reymond’s mention of Goethe alluded to his essay“The Epochs of Thought.” In this short work, Goethe dividedhistory into four ages: Poetry, Theology, Philosophy, and Prose,the last of which he judged a disaster (Goethe, 1817). DuBois-Reymond didn’t entirely share Goethe’s pessimism, perhapsbecause he equated the prosaic age with science, and perhapsbecause he felt more comfortable at the lectern. Whateverthe reason, du Bois-Reymond approached writing with a self-awareness that verged on irony. He was acutely aware of hisplace in the history of science, and his accounts of it draw fromthe language of scripture and legend. In a literal sense, du Bois-Reymond not only shifted the paradigm of neuroscience, he alsoset it.ReferencesClarke, E., and Jacyna, L. S. (1987). Nineteenth-Century Origins of NeuroscientificConcepts. Berkeley; Los Angeles; London: University of CaliforniaPress.du Bois-Reymond, E. (1848–1884). Untersuchungen über thierische Electricität.Vol. 2. Berlin: Reimer.du Bois-Reymond, E (1853). Letter to Jeannette du Bois-Reymond. Berlin:Staatsbibliothek zu Berlin, Preußischer Kulturbesitz, Haus Potsdamer Straße,Handschriftenabteilung.du Bois-Reymond, E. (1877). Letter to Jeannette du Bois-Reymond, Elberfeld:Staatsbibliothek zu Berlin, Preußischer Kulturbesitz, Haus Potsdamer Straße,Handschriftenabteilung.du Bois-Reymond, E. (1882). Letter to Jeannette du Bois-Reymond, Berlin:Staatsbibliothek zu Berlin, Preußischer Kulturbesitz, Haus Potsdamer Straße,Handschriftenabteilung.du Bois-Reymond, E. (ed.). (1912). Reden von Emil du Bois-Reymond. Leipzig:Veit.du Bois-Reymond, E. (ed.). (1918). Jugendbriefe von Emil du Bois-Reymond anEduard Hallmann. Berlin: Reimer.Finger, S. (1994). Origins of Neuroscience: A History of Explorations into BrainFunction. New York, NY: Oxford University Press.Finkelstein, G. (2013). Emil du Bois-Reymond: Neuroscience, Self, andSociety in Nineteenth-Century Germany. Cambridge; London: The MITPress.Goethe, J. W. (1817). Geistes-Epochen, nach Hermanns neuesten Mitteilungen.Über Kunst und Altherthum 1, 107–112.Gregory, F. (1977). Scientific Materialism in Nineteenth Century Germany.Dordrecht; Boston: Reidel.Heger, P. (1896–1897). Emil du Bois-Reymond. Conférence donnéeà l’Association des Étudiants en médicine de l’Université librede Bruxelles, le 26 mars 1897. Revue de l’Université de Bruxelles2, 565.Kuhn, T. S. (1961). The function of measurement in modern physical science. Isis52, 161–193. doi: 10.1086/349468McComas, A. J. (2011). Galvani’s Spark: The Story of the Nerve Impulse. New York,NY: Oxford University Press.Pera, M. (1992). The Ambiguous Frog: The Galvani-Volta Controversy onAnimal Electricity, tr. Jonathan Mandelbaum. Princeton: Princeton UniversityPress.Frontiers in Systems Neuroscience | www.frontiersin.org 3 September 2015 | Volume 9 | Article 133Finkelstein Mechanical neurosciencePiccolino, M., and Bresadola, M. (2013). Shocking Frogs: Galvani, Volta and theElectric Origins of Neuroscience. Oxford: Oxford University Press.Piccolino, M. (1998). Animal electricity and the birth of electrophysiology: thelegacy of Luigi Galvani. Brain Res. Bull. 46, 381–407. doi: 10.1016/S0361-9230(98)00026-4Thompson, W. (1889). Electrical units of measurement. Popular Lect. Address. 1,73–136.Valenstein, E. S. (2005). The War of the Soups and the Sparks: The Discovery ofNeurotransmitters and the Dispute Over How Nerves Communicate. New York,NY: Columbia University Press.Conflict of Interest Statement: The author declares that the research wasconducted in the absence of any commercial or financial relationships that couldbe construed as a potential conflict of interest.Copyright © 2015 Finkelstein. This is an open-access article distributed under theterms of the Creative Commons Attribution License (CC BY). The use, distribution orreproduction in other forums is permitted, provided the original author(s) or licensorare credited and that the original publication in this journal is cited, in accordancewith accepted academic practice. No use, distribution or reproduction is permittedwhich does not comply with these terms.Frontiers in Systems Neuroscience | www.frontiersin.org 4 September 2015 | Volume 9 | Article 133

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