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Muscles or Movements? Representation in the Nascent Brain Sciences

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Abstract

The idea that the brain is a representational organ has roots in the nineteenth century, when neurologists began drawing conclusions about what the brain represents from clinical and experimental studies. One of the earliest controversies surrounding representation in the brain was the “muscles versus movements” debate, which concerned whether the motor cortex represents complex movements or rather fractional components of movement. Prominent thinkers weighed in on each side: neurologists John Hughlings Jackson and F.M.R. Walshe in favor of complex movements, neurophysiologist Charles Sherrington and neurosurgeon Wilder Penfield in favor of movement components. This essay examines these and other brain scientists’ evolving notions of representation during the first eighty years of the muscles versus movements debate (c. 1873–1954). Although participants agreed about many of the superficial features of representation, their inferences reveal deep-seated disagreements about its inferential role. Divergent epistemological commitments stoked conflicting conceptions of what representational attributions imply and what evidence supports them.

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Notes

  1. Despite such discussions, Eccles had been discouraged from entertaining philosophical themes at the symposium. He recounts the following anecdote: “Early in the planning I was instructed by the Chancellor that ‘the Academy by its constitution has for aim to promote the study and progress of the physical, mathematical, and natural sciences and their history. Thus the discussion of philosophical questions is excluded.’ I replied, ‘…[T]o me all sciences have a philosophical basis and it is generally agreed that there is a philosophy of science which is in fact basic to all scientific investigations and discussions. Certainly when one comes to a Study Week devoted to brain and mind it is not possible to exclude relations with philosophy…” (Eccles 1965, pp. vii–viii).

  2. Mendelsohn (1987) suggested that scientific controversies are a particularly useful site for analyzing the interplay between social and conceptual factors in science. While I do here discuss professional tensions within the brain sciences, there is much interesting work yet to be done situating the debate with respect to other cultural and political currents (Pauly 1983; Harrington 1987).

  3. This section draws on Clarke and Jacyna’s (1987) Nineteenth-Century Origins of Neuroscientific Concepts, Chapters 4 and 6; Harrington’s (1987) Medicine, Mind, and the Double Brain; and Young’s (1970) Mind, Brain and Adaptation in the Nineteenth Century.

  4. Walshe (1961) traced modern representation-talk back to Müller in the 1830s. Hughlings Jackson’s contemporary, Vienna psychiatrist Theodor Meynert, also proposed a physiological theory involving distinct Vorstellungen (sometimes translated “representations”) in different cortical cells (Guenther 2015).

  5. In what remains, references to muscles proponents should accordingly be understood as shorthand, referring to those who endorse the representation of fractional movements, body parts, individual muscles, or muscle groups.

  6. Unlike most previous researchers, Penfield was also able to directly study sensory responses to stimulation, since his human patients, being only locally anesthetized, were able to verbally report what they felt. See Elder (2015) for an account of how patients acted as surveyors in Penfield’s cortical cartography project.

  7. For a different explanation of Jacksonian marches in terms of cortical representations of movement components, see Penfield and Jasper (1954, pp. 359–360).

  8. This musical metaphor has an interesting precursor in Müller’s Elements of Physiology, Vol. II: “The fibres of all the motor, cerebral and spinal nerves may be imagined as spread out in the medulla oblongata, and exposed to the influence of the will like the keys of a piano-forte” (1842, p. 934; trans. Baly). Here, in accordance with the earlier orthodoxy that motor control does not involve the cortex, the medulla oblongata (in the brainstem) is taken to be the “source of all the voluntary movements,” and the pianist is imagined to be the will, rather than other parts of the nervous system (Müller 1842, p. 934). I do not know if this image influenced the later one.

  9. Chirimuuta (2019) makes a related observation about Hughlings Jackson’s concept of representation: “unlike the computational notions prevalent today, it has none of the connotation of representations being ‘disembodied’ or abstracted from their biological basis, as software is from hardware” (p. 41). As far as I am aware, the first contributor to the muscles versus movement debate to make a clear vehicle/content distinction was Clark (1948).

  10. The possibility of recovery following brain injury was often taken to be a significant challenge for any attempt to localize functions in the brain (Harrington 1987, pp. 262–264).

  11. Even though most advocates of the representation of movements were clinical neurologists, this effort to gather experimental support for movements seems to have been led by neurophysiologist Ernst Gellhorn. With little existing scholarship on Gellhorn, it is difficult to figure out his place in our narrative. By mid-century, neurosurgeons had largely triumphed over neurologists in their struggle for professional dominance (Gavrus 2011). Perhaps the fizzling out of professional conflict removed the impetus for aligning one’s position in the muscles versus movements debate with one’s professional interests. Gellhorn’s experimental work may herald the beginning of this breakdown.

  12. In the debate’s early years, Hughlings Jackson had, in fact, made a handful of oblique comments about his use of the term “representation,” though his remarks are hardly clarifying. He claimed, for instance, that the “[m]ethod of representation and localisation are only different names for one thing” (Hughlings Jackson 1882, p. 33) and “representation and co-ordination are the same thing” (Hughlings Jackson 1884, p. 83). Presumably this means that a brain region represents a particular movement when it coordinates that movement, or the movement is localized there, though this is hardly satisfying as an explication of representation. Chirimuuta (2019) has recently offered a detailed analysis of Hughlings Jackson’s elusive notion of representation, arguing that it combines reductive and holistic strands of thought. The result, she concludes, is a uniquely “integrative” concept on which cortical representations “somehow holographically represent the entire region of the body relevant to the movement…while at the same time co-ordinating or controlling more local parts” (Chirimuuta 2019, p. 41).

  13. Walshe also declared that representations can be distributed, or “spread out widely throughout the motor cortex” (1943, p. 111). For decades, participants in the debate had closely associated representation with localization, assuming that representations occupy restricted areas of the cortex. Walshe drove a wedge between representation and localization by claiming that cortical representations are “widely localized” (1943, p. 111).

  14. McMullin (1987) suggests that a difference between scientific and moral controversies is that there is usually consensus about the proper means of resolving the former but not the latter. In scientific controversies, unlike moral ones, participants “agree (at least in broad outline) on what would count as relevant considerations on either side of the case” (p. 55). The muscles versus movements debate shows this is unreasonably optimistic. Because participants did not share a common understanding of representation, there was little agreement about what would constitute decisive evidence for one side or the other.

  15. This brief discussion of inferentialism elides many nuances. For one thing, the inferences that Brandom holds determine a concept’s meaning are correct inferences: “the ones people ought to make, rather than the ones they are actually disposed to make” (2010, p. 163). For another, Brandom recognizes that the inferential significance of a concept depends on auxiliary hypotheses, or what he calls “collateral commitments” (1994, p. 92). Perhaps the disagreements between brain scientists discussed here can be traced back to differing collateral commitments. An objector might claim that the meaning of “representation” was shared among scientists, even if they were led to endorse different inferences by different auxiliaries. A more careful articulation of the inferentialist argument suggested here would need to address these complexities.

  16. Criticism of the concept of representation can be found in other literatures that are more or less related to the present discussion of representation in the brain. For instance, science studies scholars have debated the utility of the notion of representation (Lynch and Woolgar 1990; Lynch 1994), and the “representation wars” in cognitive science have pitted representationalist against dynamicist models of cognitive processes (Constant et al. 2021).

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Acknowledgements

My thanks to attendees of FSU’s Philosophical Psychology Group, the 2021 APA Pacific, 2021 PSA, and 2021 Deep South Philosophy and Neuroscience Workgroup for their feedback. Special thanks to Dan Weiskopf for his APA commentary; to Edouard Machery for pointing me to Rheinberger’s work on conceptual mutability; to two anonymous reviewers for JHB for their very constructive comments; and to Liam Kofi Bright, Ron Doel, Jack Justus, Jacob Neal, and Shivam Patel for feedback on earlier drafts of the paper.

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Ward, Z.B. Muscles or Movements? Representation in the Nascent Brain Sciences. J Hist Biol 56, 5–34 (2023). https://doi.org/10.1007/s10739-023-09703-1

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