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Life of the Cell Philosophy & History of Cell Research A workshop at Egenis, University of Exeter April 23rd & 24th, 2009 •William Bechtel •Thomas Cavalier-Smith •Ohad Parnes •Andrew Reynolds Life of the cell: Philosophy & history of cell research On April 23rd and 24th, 2009, at Egenis, University of Exeter, UK, a number of issues related to cells and cell biology will be investigated from historical, philosophical and scientific perspectives. Topics include: •The cell as the epistemic and ontological unit of life •The evolution of cells and modes of cellular organization • Multicellular and symbiotic cell collectives in relation to cellular autonomy • Connections between cell biology and contemporary systems biology • Plant, fungi and insect cells that call standard cell theory into question A pre-workshop reading list is available on the workshop website: www.genomicsnetwork.ac.uk/ egenis/events/workshops/title,3800,en.html Keynote speakers William Bechtel (Philosophy, University of California, San Diego): The Cell: Locus or object of inquiry? Thomas Cavalier-Smith (Zoology, University of Oxford): The evolution of cells Ohad Parnes (History, Central European University, Budapest): Agents as cells: Conceptual and historical perspectives Andrew Reynolds (Philosophy and Religious Studies, Cape Breton University, Nova Scotia): The redoubtable cell Speakers (Egenis, University of Exeter) John Bryant: Plant form: Cell differentiation with and without walls Steve Hughes: Plant form: Cell differentiation with and without walls Commentators Sabine Brauckmann (Science Centre, Tartu University, Estonia) John Dupré (Egenis, University of Exeter) Pierre-Olivier Méthot (Egenis, University of Exeter) Staffan Müller-Wille (Egenis, University of Exeter) Dan Nicholson (Egenis, University of Exeter) Maureen O’Malley (Egenis, University of Exeter) Contact Saira Kidangan, Egenis Secretary Byrne House, University of Exeter St Germans Road, Exeter, EX4 4PJ, UK Email: [email protected] Phone: +44 (0)1392 269140 Workshop programme Thursday, April 23rd 8.30-9.00 Coffee 9.00-9.30 Welcome and introduction John Dupré & Staffan Müller-Wille 9.30-10.30 William Bechtel The Cell: Locus or Object of Inquiry? 10.30-11.00 Coffee break 10.45-11.15 Dan Nicholson Commentary on Bechtel 11.15-11.45 John Dupré Commentary on Bechtel 11.45-12.00 General discussion 12.00-1.30 Lunch 1.30-2.30 2.30-3.00 3.00-3.30 3.30-4.30 4.30-5.00 5.00-5.15 Andrew Reynolds The redoubtable cell Sabine Brauckmann Commentary on Reynolds Coffee break Ohad Parnes Agents as cells: Conceptual & historical perspectives Staffan Müller-Wille Commentary on Parnes General discussion Evening Drinks and dinner Friday, April 24th 8.30-9.00 Coffee 9.00-10.00 Thomas Cavalier-Smith The evolution of cells: Real history is messy and non-Platonic 10.00-10.15 Andrew Reynolds Commentary on Cavalier-Smith 10.15-10.30 Maureen O’Malley Commentary on Cavalier-Smith 10.30-10.45 General discussion 10.45-11.15 Coffee break 11.15-12.15 John Bryant & Steve Hughes (joint presentation) Plant form: Cell differentiation with and without walls 12.15-12.30 General discussion (15 minutes) 12.30-1.30 Summary and roundtable Led by Pierre-Olivier Méthot & Dan Nicholson 1.30 Lunch Life of the Cell: Philosophy & History of Cell Research A pre-workshop reading list is available on the workshop website: www.genomicsnetwork.ac.uk/egenis/events/workshops/title,3800,en.html Titles and abstracts The Cell: Locus or object of inquiry? WILLIAM BECHTEL Department of Philosophy and Interdisciplinary Programs in Cognitive Science and Science Studies University of California, San Diego The adoption of the name “cell biology” in the middle years of the 20th century for a field of inquiry suggested the privileging of cells as objects of inquiry. The term was chosen to supplant the older term “cytology” to reflect the productive confluence of techniques—electron microscopy and cell fractionation—that enabled an integrated investigation into cell structure and function. In fact, however, in important respects the cell was soon left behind as research decomposed cells into organelles and systems of enzymes and focused on these. For some theorists, such reductionistic inquiry is synonymous with progress. But is the cell now just a locus of inquiry, no longer important as an object in its own right? The growing appeals to systems in biology suggest its continued importance as an object of inquiry. The decomposition into parts and operations is being complemented by recomposition in which the interactions between components, often dynamic, become the focus of inquiry. For some purposes researchers are recognizing the need to situate the cell in its environment to understand operations within it. For illustration, I will appeal to research on circadian clocks in which the enormous success of research decomposing individual cells that maintain circadian oscillations into genes and proteins in the 1990s is complemented by research recomposing systems within cells and research situating individual cells in larger networks. The cell remains not just a locus but also an object of inquiry. Life of the Cell Titles and abstracts (continued) The evolution of cells: Real history is messy and non-Platonic THOMAS CAVALIER-SMITH Department of Zoology, University of Oxford, South Parks Road, Oxford OX1 3PS, United Kingdom I shall discuss four main questions. First, the historical role of studies of protozoa and unicellular algae in the early history of the cell theory (mainly first half of 19th century). This focuses on recognition of the nucleus and its sphere of influence and of cell lineages as key units for much of biology. Second, the long drawn out, often confused, recognition that bacteria are structurally radically different from, yet ancestral to nucleated cells. Some have held that we should therefore not treat bacteria as cells. But we usually do. To unify both types of cell into a single concept we must regard a cell as a holistic assemblage that grows and divides as a result of the integrated activity of five kinds of entity: bounding (and optionally internal) membrane(s); genes constituting one or more chromosomes (DNA molecules); a cell skeleton; catalysts; and a pervasive watery milieu containing mineral ions and small organic molecules. Third, I shall give some microbial examples of the fuzziness of the distinction between cells and syncytia and the non-text-book nature of many body plans. Fourth, I shall discuss the permanent evolutionary merger of distinct cells to form radically new chimaeric supercells, with special reference to the question when an obligately endosymbiotic foreign cell makes the transition to an integrated cell organelle, as in the origin of mitochondria from purple bacteria, of chloroplasts from cyanobacteria, and of the exceedingly complex chromalveolate cells that arose when a biciliate protozoan enslaved and grossly modified a red algal eukaryotic cell. In so doing I shall touch on a variety of issues of possible historical or philosophic interest, including the idea of membrane heredity, often arguing that real history and biology are messy and often excessively oversimplified for didactic or other reasons. Yet the purpose of science and philosophy is to oversimplify boldly but quasi-realistically. Life of the Cell Titles and abstracts (continued) Agents as cells: Conceptual and historical perspectives OHAD S. PARNES Associate Professor History Department Central European University Budapest And: Head of Research Group: History of Heredity Zentrum für Literatur- und Kulturforschung Berlin I have argued elsewhere that contrary to common belief the ‘discovery’ of the cell theory by Theodor Schwann in 1838 was not so much a matter of better microscopy, but a direct result of an attempt to provide a mechanisticlike explanation of physiological processes in terms of their specific agents – namely cells. At least since the beginning of the twentieth century, cells seem to have lost their special role as agents of physiological or pathological processes. In genetics, the agency of heredity has been located first in chromosomes, then in genes and later in the molecular structure of DNA. In pathology, the focus has similarly shifted from cells to their components – e.g. the interaction with viral DNA/RNA in infection. At the same time, other areas of biomedicine have favoured explanatory schemes in which the cell is but an actor within a system, and where the agency is ascribed to the behaviour of the system as a whole, not to the activity of single cells (or their components) – here the most notable examples are from neurology and immunology. And yet, exactly hundred and seventy years after the publication of Theodor Schwann’s work, there seems to be no way of relinquishing cells in biomedical explanations. My talk will be an attempt to address this enigmatic role of the cell in modern biomedicine: Is the cell in the 21st century merely a morphological unit? To what extent has it retained its original role as an agent of life and disease? Is there a place for cells also in explanatory schemes where agency has been exchanged for reactivity? And finally: Does it make sense to talk about one cell concept in modern biomedicine or is it the case that different disciplines are employing different notions of cells (e.g. immunological cells, neurological cells, microbiological cells, etc.)? Life of the Cell: Titles and abstracts (continued) The redoubtable cell ANDREW REYNOLDS Department of Philosophy and Religious Studies Cape Breton University From its first introduction the cell theory – the thesis that all life is situated within a fundamental morphological and physiological unit, typically of microscopic size – has faced criticism. Yet despite frequent charges of its inadequacy the cell theory has (to borrow a line from the French philosopher Gilson) continued to bury its undertakers. Why? What is it about the cell concept that provides it with this redoubtable quality? (The urge to make a pun here is irresistible, since redoubtable could also be construed to mean always subject to doubt.) My talk will look at the history of criticisms of the cell doctrine in an attempt to identify why it is that despite its oft-noted shortcomings the cell concept continues to enjoy such vitality. Plant form: Cell differentiation with and without walls JOHN BRYANT and STEVE HUGHES Biosciences & Egenis, University of Exeter It is generally recognised that cell migration is not a feature of plant development and that plant form is forged by anisotropic cell expansion coupled to the reorientation of planes of division well in advance of mitosis (Lloyd and Chan 2005) and also by asymmetric divisions. For a more complete picture however, we need to add to this recipe the more subtle developmental themes of apoptosis, or programmed cell death (Rogers, 2005), conductive tissues of intermediate cellular status (Holbrook et al., 2002; van Bel, 2003) and the nesting of cells within cells in the sporophytegametophyte transition. Using examples of phloem and xylem cells (with walls), aerenchyma , the egg apparatus, and syncytial endosperm (without walls) we will examine these developmental themes against the proposal of Baluka et al., (2004) for a partial disaggregation of cellular space into nuclear and peripheral components (cell bodies). In the case of syncytial endosperm it seems most interesting that a regular zonal differentiation of mitotic rhythm can occur among nuclei, setting up the endosperm structural axis in advance of cellularization of the space itself (Boisnard-Lorig et al., 2001). We therefore finish with a question - When is a cell not a cell? and with its corollary - When is an entity that is 'not a cell' a cell?