(Recipient of the 2020 Everett Mendelsohn Prize.) This article revisits the development of the protoplasm concept as it originally arose from critiques of the cell theory, and examines how the term “protoplasm” transformed from a botanical term of art in the 1840s to the so-called “living substance” and “the physical basis of life” two decades later. I show that there were two major shifts in biological materialism that needed to occur before protoplasm theory could be elevated to have equal status (...) with cell theory in the nineteenth century. First, I argue that biologists had to accept that life could inhere in matter alone, regardless of form. Second, I argue that in the 1840s, ideas of what formless, biological matter was capable of dramatically changed: going from a “coagulation paradigm” that had existed since Theophrastus, to a more robust conception of matter that was itself capable of movement and self-maintenance. In addition to revisiting Schleiden and Schwann’s original writings on cell theory, this article looks especially closely at Hugo von Mohl’s definition of the protoplasm concept in 1846, how it differed from his primordial utricle theory of cell structure two years earlier. This article draws on Lakoff and Johnson’s theory of “ontological metaphors” to show that the cell, primordial utricle, and protoplasm can be understood as material container, object, and substance, and that these overlapping distinctions help explain the chaotic and confusing early history of cell theory. (shrink)

When discussing the changing sense of reality around 1900 in the cultural arts the lexicon of early modernism reigns supreme. This essay contends that a critical condition for the possibility of many of the turn of the century modernist movements in the arts can be found in exchange of instruments, concepts, and media of representation between the sciences and the arts. One route of interaction came through physiological aesthetics, the attempt to ‘elucidate physiologically the nature of our Aesthetic feelings’ and (...) explain how works of art achieve their effects. Physiological aesthetics provided the terms for new formalist languages of art and criticism, and in some instances suggested optimistic, even utopian, possibilities for art to remake human individuals and societies.Keywords: Physiology; Psychology; Evolution; Aesthetics; Modernism; Art history. (shrink)

This essay aims to shed new light on the relations between physiology and psychology in the late nineteenth and early twentieth centuries by focusing on the use of unicellular organisms as research objects during that period. Within the frameworks of evolutionism and monism advocated by Ernst Haeckel, protozoa were perceived as objects situated at the borders between organism and cell and individual and society. Scholars such as Max Verworn, Alfred Binet, and Herbert Spencer Jennings were provoked by these organisms to (...) undertake experimental investigations situated between general physiology and psychology that differed from the physiological psychology advocated by Wilhelm Wundt. Some of these investigations sought to locate psychological properties in the molecular structure of protoplasm; others stressed the existence of organic and psychological individuality in protozoa. In the following decades, leading philosophers such as Friedrich Nietzsche, Charles Sanders Peirce, and Henri Bergson, as well as psychological researchers like Sigmund Freud, integrated the results of these investigations into their reflections on such problems as the nature of the will, the structure of the ego, and the holistic nature of the reactions of organisms to their environment. (shrink)

In 1853, the young Thomas Henry Huxley published a long review of German cell theory in which he roundly criticized the basic tenets of the Schleiden-Schwann model of the cell. Although historians of cytology have dismissed Huxley's criticism as based on an erroneous interpretation of cell physiology, the review is better understood as a contribution to embryology. "The Cell-theory" presents Huxley's "epigenetic" interpretation of histological organization emerging from changes in the protoplasm to replace the "preformationist" cell theory of Schleiden and (...) Schwann (as modified by Albert von Kölliker), which posited the nucleus as the seat of organic vitality. Huxley's views influenced a number of British biologists, who continued to oppose German cell theory well into the twentieth century. Yet Huxley was pivotal in introducing the new German program of "scientific zoology" to Britain in the early 1850s, championing its empiricist methodology as a means to enact broad disciplinary and institutional reforms in British natural history. (shrink)

In the nineteenth century protozoology and early cell biology intersected through the nexus of Darwin's theory of evolution. As single-celled organisms, amoebae offered an attractive focus of study for researchers seeking evolutionary relationships between the cells of humans and other animals, and their primitive appearance made them a favourite model for the ancient ancestor of all living things. Their resemblance to human and other metazoan cells made them popular objects of study among morphologists, physiologists, and even those investigating animal behaviour. (...) The amoeba became the exemplar of the new protoplasmic cell concept of mid-century and because its apparent simplicity made it widely generalizable it became a popular subject in a breadth of experimental investigations and theoretical speculations. It was able to do this because "the amoeba" denotes not a particular organism, but a general type of behaviour common to the cells of a range of protozoa, simple plants and higher animals. Its status as an exemplary cell also rested upon auxiliary philosophical assumptions about what constitutes a primitive characteristic and the thesis that evolution is a progressive development of order from chaos. (shrink)

Between 1900 and 1930 colloid chemistry, a branch of physical chemistry, gained crucial importance for the understanding of vital phenomena. To many it seemed that the properties of colloids would differ from those of ordinary matter, paralleling the specific properties of protoplasm, the living substance . The application of theoretical concepts and experimental models of colloid chemistry to biological problems shaped Biocolloidology as a new research program, which appeared promising for the exploration of organic processes such as mitotic cell division, (...) growth, muscle contraction and movement. Biocolloidology could simultaneously absorb the quest for a physical-chemical explanation of life and the (post)-romantic notion of organic nature as a container of creative and vital forces that was central to life-philosophy. At least until the late 1920s, it is argued, the merging of different scientific and philosophical metaphors made Biocolloidology and the concept of protoplasm highly appealing in a broad cultural context. (shrink)

What is the relation between things and theories, the material world and its scientific representations? This is a staple philosophical problem that rarely counts as historically legitimate or fruitful. In the following dialogue, the interlocutors do not argue for or against realism. Instead, they explore changing relations between theories and things, between contested objects of knowledge and less contested, more everyday things. Widely seen as the life sciences' first general theory, the cell theory underwent dramatic changes during the nineteenth century. (...) The dialogue establishes that each successive version of the cell theory was formulated -- each identity of the object cell was formed -- around a different material : cork, cartilage, eggs in cleavage, muscle. Such things thus serve as exemplary materials, in ways not described by standard concepts like induction, theory -testing, theory -laden observation, and construction. Still, how can theories and perspective possibly be honed on things if these are apprehended differently by different observers according to their interests, training, culture, or indeed theories? The second part of the dialogue addresses this problem, partly through the verbal and visual schemata that were used by nineteenth-century microscopists and that are comparable to schemata in the visual arts. The third part of the dialogue considers the exemplary materials as a historical sequence, itself needing explanation. Theoretical change devolved partly from wider histories and geographies of the prevalence, availability, or scientific and cultural status of materials such as plants, animals, and muscle. (shrink)

Heredity has been dismissed as an insignificant object in Claude Bernard’s physiology, and the topic is usually ignored by historians. Yet, thirty years ago, Jean Gayon demonstrated that Bernard did elaborate on the subject. The present paper aims at reassessing the issue of heredity in Claude Bernard’s project of a “general physiology”. My first claim is that Bernard’s interest in heredity was linked to his ambitious goal of redefining general physiology in relation to morphology. In 1867, not only was morphology (...) included within experimental physiology, but it also theoretically grounded physiological investigations. By 1878, morphology and physiology were considered as completely independent sciences, and only the latter was perceived as suitable to experimentation. My second claim is that this reversal reflected the existence of two opposite attitudes towards heredity. In the late 1860s, Bernard was convinced that heredity would soon be accessible to experimental manipulation and that new species would be produced in the laboratory exactly like organic chemistry succeeded to do for raw bodies. Ten years later, he ascertained that this was impossible. My third claim is that Bernard was epistemologically ill-equipped to address the issue of heredity. Bernard was strongly committed to a general reasoning scheme that acknowledged only three categories: determining conditions, constant laws and phenomena. This scheme was a key factor in his successes as a physiologist who was able to capture new mechanisms in living bodies. Nonetheless, it also prevented him from understanding how time and history could be endowed with a causal action that cannot be reduced to timeless parameters. (shrink)

The rise of the mechanistic worldview in the seventeenth century had a major impact on views of biological generation. Many seventeenth century naturalists rejected the old animist thesis. However, the alternative view of gradual mechanistic formation in embryology didn’t convince either. How to articulate the peculiarity of life? Researchers in the seventeenth century proposed both “animist” and mechanistic theories of life. In the eighteenth century again a controversy in biology arose regarding the explanation of generation. Some adhered to the view (...) that life is a physical property of matter (e.g. Buffon), others saw living entities as the result of the development of pre-existing germs (e.g. Bonnet). Naturalists, lacked a convincing account that could guide their research. In interaction with leading naturalists of his time Immanuel Kant articulated an approach to explaining generation. Kant’s account, delineated in his Kritik der Urteilskraft (Critique of the power of judgment) (1790), is a combination of Newtonian non-reductionist mechanism in explanation, and a concept of natural end comparable to Stahl’s formal conception of organic bodies. It consists of two claims: a) in biology only mechanical explanation is explanatory, and b) living entities contain some original organisation, which is mechanically unexplainable. In the nineteenth century this approach influenced naturalists as Müller, Virchow, and Von Baer, in their physiological research. Dissatisfied with a sheer mechanistic or, on the other hand, a sheer teleological approach, they appreciated the Kantian account of mechanical explanation of natural ends. In Germany, in the second halve of the nineteenth century, Ernst Haeckel reopened the debate about abiogenesis, which still continuous. (shrink)

The question concerning the meaning of life is important, but it immediately confronts the present authors with insurmountable obstacles from a philosophical standpoint, as it would require us to define not only what we hold to be life, but what we hold to be meaning in addition, requiring us to do both in a properly researched context. We unconditionally surrender to that challenge. Instead, we offer a vernacular, armchair approach to life’s origin and meaning, with some layman’s thoughts on the (...) meaning of origins as viewed from the biologist’s standpoint. One can observe that biologists generally approach the concept of biological meaning in the context of evolution. This is the basis for the broad resonance behind Dobzhansky’s appraisal that “Nothing in biology makes sense except in the light of evolution”. Biologists try to understand living things in the historical context of how they arose, without giving much thought to the definition of what life or living things are, which for a biologist is usually not an interesting question in the practical context of daily dealings with organisms. Do humans generally understand life’s meaning in the context of history? If we consider the problem of life’s origin, the question of what constitutes a living thing becomes somewhat more acute for the biologist, though not more answerable, because it is inescapable that there was a time when there were no organisms on Earth, followed by a time when there were, the latter time having persisted in continuity to the present. This raises the question of where, in that transition, chemicals on Earth became alive, requiring, in turn, a set of premises for how life arose in order to conceptualize the problem in relation to organisms we know today, including ourselves, which brings us to the point of this paper: In the same way that cultural narratives for origins always start with a setting, scientific narratives for origins also always start with a setting, a place on Earth or elsewhere where we can imagine what happened for the sake of structuring both the problem and the narrative for its solution. This raises the question of whether scientific origins settings convey meaning to humans in that they suggest to us from what kind of place and what kinds of chemicals we are descended, that is, to which inanimate things we are most closely related. (shrink)

Since the early nineteenth century a membrane or wall has been central to the cell’s identity as the elementary unit of life. Yet the literally and metaphorically marginal status of the cell membrane made it the site of clashes over the definition of life and the proper way to study it. In this article I show how the modern cell membrane was conceived of by analogy to the first “artificial cell,” invented in 1864 by the chemist Moritz Traube (1826–1894), and (...) reimagined by the plant physiologist Wilhelm Pfeffer (1845–1920) as a precision osmometer. Pfeffer’s artificial cell osmometer became the conceptual and empirical basis for the law of dilute solutions in physical chemistry, but his use of an artificial analogue to theorize the existence of the plasma membrane as distinct from the cell wall prompted debate over whether biology ought to be more closely unified with the physical sciences, or whether it must remain independent as the science of life. By examining how the histories of plant physiology and physical chemistry intertwined through the artificial cell, I argue that modern biology relocated vitality from protoplasmic living matter to nonliving chemical substances—or, in broader cultural terms, that the disenchantment of life was accompanied by the (re)enchantment of ordinary matter. (shrink)

This essay details a historical crossroad in biochemistry and microbiology in which penicillin was a co-agent. I narrate the trajectory of the bacterial cell wall as the precise target for antibiotic action. As a strategic object of research, the bacterial cell wall remained at the core of experimental practices, scientific narratives and research funding appeals throughout the antibiotic era. The research laboratory was dedicated to the search for new antibiotics while remaining the site at which the mode of action of (...) this new substance was investigated. This combination of circumstances made the bacterial wall an ontology in transit. As invisible as the bacterial wall was for clinical purposes, in the biological laboratory, cellular meaning in regard to the action of penicillin made the bacterial wall visible within both microbiology and biochemistry. As a border to be crossed, some components of the bacterial cell wall and the biochemical destruction produced by penicillin became known during the 1950s and 1960s. The cell wall was constructed piece by piece in a transatlantic circulation of methods, names, and images of the shape of the wall itself. From 1955 onwards, microbiologists and biochemists mobilized new names and associated conceptual meanings. The composition of this thin and rigid layer would account for its shape, growth and destruction. This paper presents a history of biochemical morphology: a chemistry of shape – the shape of bacteria, as provided by its wall – that accounted for biology, for life itself. While penicillin was being established as an industrially-manufactured object, it remained a scientific tool within the research laboratory, contributing to the circulation of further scientific objects. (shrink)

This paper explores Charles Darwin’s hypothesis of pangenesis through a popular and professional reception history. First published in The Variation of Animals and Plants under Domestication (1868), pangenesis stated that inheritance can be explained by sub-cellular “gemmules” which aggregated in the sexual organs during intercourse. Pangenesis thereby accounted for the seemingly arbitrary absence and presence of traits in offspring while also clarifying some botanical and invertebrates’ limb regeneration abilities. I argue that critics largely interpreted Variation as an extension of On (...) the Origin of Species by Means of Natural Selection (1859), while pangenesis was an extension of natural selection. Contrary to claims that pangenesis was divorced from natural selection by its reliance on the inheritance of acquired characters, pangenesis’s mid nineteenth-century reception suggests that Darwin’s hypothesis responded directly to selection’s critics. Using Variation’s several editions, periodical reviews, and personal correspondence I assess pangenesis popularly, professionally, and biographically to better understand Variation’s impact on 1860s and 70s British evolutionism and inheritance. -/- . (shrink)

What are cells? How are they related to each other and to the organism as a whole? These questions have exercised biology since Schleiden and Schwann (1838–1839) first proposed cells as the key units of structure and function of all living things. But how do we try to understand them? Through new technologies like the achromatic microscope and the electron microscope. But just as importantly, through the metaphors our culture has made available to biologists in different periods and places. These (...) two new volumes provide interesting history and philosophy of the development of cell biology. Reynolds surveys the field's changing conceptual structure by examining the varied panoply of changing metaphors used to conceptualize and explain cells – from cells as empty boxes, as building blocks, to individual organisms, to chemical factories, and through many succeeding metaphors up to one with great currency today: cells as social creatures in communication with others in their community. There is some of this approach in theVisionsedited collection as well. But this collection also includes rich material on the technologies used to visualize cells and their dialectical relationship with the epistemology of the emerging distinct discipline of cell biology. This volume centres on, but is not limited to, ‘reflections inspired by [E.V.] Cowdry's [1924 volume]General Cytology’; it benefits from a conference on the Cowdry volume as well as a 2011 Marine Biological Lab/Arizona State University workshop on the history of cell biology. (shrink)