The redoubtable cell Andrew Reynolds Department of Philosophy and Religious Studies, Cape Breton University, 1250 Grand Lake Road, Sydney, NS B1P 6L2, Canada article info Keywords: Cell theory Cell standpoint Organism standpoint abstract The cell theory—the thesis that all life is made up of one or more cells, the fundamental structural and physiological unit—is one of the most celebrated achievements of modern biological science. And yet from its very inception in the nineteenth century it has faced repeated criticism from some biologists. Why do some continue to criticize the cell theory, and how has it managed nevertheless to keep burying its undertakers? The answers to these questions reveal the complex nature of the cell theory and the cell concept on which it is based. Like other scientific ‘laws’, the assertion that all living things are made of cells purchases its universality at the expense of abstraction. If, however, this law is regarded merely as a widely applicable empirical generalization with notable exceptions, it still remains too important to discard. Debate about whether the cell or the organism standpoint provides the more correct account of anatomical, physiological, and developmental facts illustrates the tension between our attempts to express the truth about reality in conceptual terms conducive to a unified human understanding. Ó 2010 Elsevier Ltd. All rights reserved. When citing this paper, please use the full journal title Studies in History and Philosophy of Biological and Biomedical Sciences Seek simplicity and distrust it. (A. N. Whitehead) 1 1. Introduction The cell theory has been said to stand next to Darwin’s theory of evolution as one of the most significant achievements of modern biological science (Wilson, 1906, p. 1; Chubb, 1911, p. 710; Mazzarello, 1999, p. E15; Nurse, 2000, p. 1711). And yet the the- ory of the cellular basis of life has faced stern criticism from biolo- gists who have disputed the adequacy of the cell doctrine and the cell concept on which it is built. Many of these critics, especially in the nineteenth and early twentieth centuries, characterized the cell concept as a Baconian idol responsible for misguiding the direction of scientific investigation. There are still some critics today who in- sist it must be rejected or at least seriously revised (for example, Kaplan & Hagemann, 1991; Sitte, 1992; Baluška et al., 2004; Mavrakis et al., 2009). And yet despite this repeated criticism the cell theory has continued to bury its undertakers. Why? What is it about the cell theory that makes it such a redoubtable opponent? I am using the word redoubtable here as a pun, in the sense that saying of something that it is redoubtable could also be construed to mean that it is always subject to doubt. The pun is more than just playful, for, as I will argue, it helps to make an important point about the complex nature of the cell theory. The history of the cell theory is inseparable, naturally enough, from the history of the cell concept; and the cell concept has been molded, revised and refashioned through the use of multiple met- aphors. The very concept of the cell as coined by Robert Hooke (1635–1703) in 1665 is of course based on the metaphor of an empty room enclosed within rigid walls (Hooke, 2007 [1665]). This provided early investigators with a helpful search image as they inspected various tissues. In the nineteenth and early twen- tieth centuries cells were frequently referred to as the Bausteine or building stones out of which the animal or plant body is con- structed. At the same time it was popular practice to refer to the cell as an elementary organism and to the whole plant or ani- mal as a composite ‘cell-state’ (Reynolds, 2007a, 2008b). This per- spective emphasized plant and animal cells as autonomous vital units that divide among themselves the physiological labour of the organismal economy. 1369-8486/$ - see front matter Ó 2010 Elsevier Ltd. All rights reserved. doi:10.1016/j.shpsc.2010.07.011 E-mail address: andrew_reynolds@cbu.ca 1 Quoted in Gannon (2007), p. 705, from Whitehead (1955), p. 163. Studies in History and Philosophy of Biological and Biomedical Sciences 41 (2010) 194–201 Contents lists available at ScienceDirect Studies in History and Philosophy of Biological and Biomedical Sciences journal homepage: www.elsevier.com/locate/shpsc