Supervenience is a relationship which has been used recently to explain the physical determination of biological phenomena despite resistance to reduction. Supervenience, however, is plagued by ambiguities which weaken its explanatory value and obscure some interesting aspects of reduction in biology. Although I suspect that similar considerations affect the use of supervenience in ethics and the philosophy of mind, I don’t intend anything I have to say here to apply outside of the physical and biological cases I consider.The main point (...) of this paper is that there is a property of biological systems which makes it both misleading and inappropriate to reduce central biological phenomena to the properties of underlying components. Despite this, reductive explanation has been a major source of innovation in biological theory. The apparent tension can be resolved if underlying properties are explanatorily relevant to the higher level phenomena even though the latter are not strictly reducible to the former. Supervenience, I will argue, is not robust enough to deny reduction while supporting explanatory relevance. (shrink)

Darwin's ideas on variation, heredity, and development differ significantly from twentieth-century views. First, Darwin held that environmental changes, acting either on the reproductive organs or the body, were necessary to generate variation. Second, heredity was a developmental, not a transmissional, process; variation was a change in the developmental process of change. An analysis of Darwin's elaboration and modification of these two positions from his early notebooks (1836-1844) to the last edition of the /Variation of Animals and Plants Under Domestication/ (1875) (...) complements previous Darwin scholarship on these issues. Included in this analysis is a description of the way Darwin employed the distinction between transmission and development, as well as the conceptual relationship he saw between heredity and variation. This paper is part of a larger project comparing commitments regarding variation during the latter half of the nineteenth century. (shrink)

August Weismann is famous for having argued against the inheritance of acquired characters. However, an analysis of his work indicates that Weismann always held that changes in external conditions, acting during development, were the necessary causes of variation in the hereditary material. For much of his career he held that acquired germ-plasm variation was inherited. An irony, which is in tension with much of the standard twentieth-century history of biology, thus exists – Weismann was not a Weismannian. I distinguish three (...) claims regarding the germ-plasm: (1) its continuity, (2) its morphological sequestration, and (3) its variational sequestration. With respect to changes in Weismann’s views on the cause of variation, I divide his career into four stages. For each stage I analyze his beliefs on the relative importance of changes in external conditions and sexual reproduction as causes of variation in the hereditary material. Weismann believed, and Weismannism denies, that variation, heredity, and development were deeply intertwined processes. This article is part of a larger project comparing commitments regarding variation during the latter half of the nineteenth century. (shrink)

From Spermatic Animalcules to Sperm Cells: The Reconceptualization of Generation in the 19th Century. At the end of the 18th and still at the beginning of the 19th century most naturalists considered spermatic animalcules to be parasites of the seminal fluid that played no role in procreation. This view was progressively questioned by 19th century physiologists. They gradually redefined the spermatic animals as (cellular) products of the male organism, as agents of fertilization and bearers of the male heredity material. This (...) article discusses this change from two different perspectives: on a microhistorical level, it analyzes the experimental research of the naturalist Lazzaro Spallanzani (1729–1799) and of the physiologist Albert Kölliker (1817–1905) in order to show how spermatozoa were turned into a new epistemic object of biology – the sperm cell. Further, it asks how the role of the reconceptualization of spermatic animalcules affected the long‐term transformations that gave rise of our modern understanding of heredity, generation and the sexed body. By combining these two perspectives, the article aims to connect historiographies that are often kept separate: the macrohistorical narratives about gender and the body in the modern age and the microhistorical studies of biomedical practices and objects. (shrink)

Cytogenetics was born of the confluence of two so-far independent fields: cytological studies and breeding studies, which merged through the identification of genes with chromosomes. In this paper I argue that genes were introduced as functional entities. Functional explanations are presented here as a subclass of inferences to the best explanation and I argue that abductive arguments do not offer conclusive proof for the existence of the entities postulated through them. However, functional explanations usually follow a scheme laid out by (...) Fodor (1968): there is a first phase where hypothetical entities are postulated and individuated through their effects; there is a second phase where the physical structures responsible for these effects are sought. I analyze the development of both phases in the construction of the chromosome theory of Mendelian inheritance. I will argue that first-phase theories are important to set conditions of identification for the functions played by a certain structure in a given containing system. (shrink)

T. H. Morgan (1866–1945), the founder of the Drosophila research group in genetics that established the chromosome theory of Mendelian inheritance, has been described as a radical empiricist in the historical literature. His empiricism, furthermore, is supposed to have prejudiced him against certain scientific conclusions. This paper aims to show two things: first, that the sense in which the term empiricism has been used by scholars is too weak to be illuminating. It is necessary to distinguish between empiricism as an (...) epistemological position and the so-called methodological empiricism. I will argue that the way the latter has been presented cannot distinguish an empiricist methodology from a non-empiricist one. Second, I will show that T. H. Morgan was not an epistemological empiricist as this term is usually defined in philosophy. The reason is that he believed in the existence of genes as material entities when they were unobservable entities when they were unobservable entities introduced to account for the phenotypic ratios found in breeding experiments. These two points, of course, are interrelated. If we were to water down the meaning of empiricis, perhaps we could call Morgan an empiricist. But then we would also fail to distinguish empiricism from realism. (shrink)

Early in his career Thomas Hunt Morgan was interested in embryology and dedicated his research to studying organisms that could regenerate. Widely regarded as a regeneration expert, Morgan was invited to deliver a series of lectures on the topic that he developed into a book, Regeneration. In addition to presenting experimental work that he had conducted and supervised, Morgan also synthesized and critiqued a great deal of work by his peers and predecessors. This essay probes into the history of regeneration (...) studies by looking in depth at Regeneration and evaluating Morgan’s contribution. Although famous for his work with fruit fly genetics, studying Regeneration illuminates Morgan’s earlier scientific approach which emphasized the importance of studying a diversity of organisms. Surveying a broad range of regenerative phenomena allowed Morgan to institute a standard scientific terminology that continues to inform regeneration studies today. Most importantly, Morgan argued that regeneration was a fundamental aspect of the growth process and therefore should be accounted for within developmental theory. Establishing important similarities between regeneration and development allowed Morgan to make the case that regeneration could act as a model of development. The nature of the relationship between embryogenesis and regeneration remains an active area of research. (shrink)

In earlier work I have argued that the most substantial threat to scientific realism arises from the problem of unconceived alternatives: the repeated failure of past scientists and scientific communities to conceive of alternatives to extant scientific theories, even when such alternatives were both (1) well confirmed by the evidence available at the time and (2) sufficiently scientifically serious as to be later embraced by actual scientific communities. In this paper I explore Charles Darwin's development and defense of his 'pangenesis' (...) theory of inheritance and conclude that this particular historical example offers impressive support for the challenge posed to realism by this problem of unconceived alternatives. (shrink)

The idea of ‘reversion’ or ‘atavism’ has a peculiar history. For many authors in the latenineteenth and early-twentieth centuries – including Darwin, Galton, Pearson, Weismann, and Spencer, among others – reversion was one of the central phenomena which a theory of heredity ought to explain. By only a few decades later, however, Fisher and others could look back upon reversion as a historical curiosity, a non-problem, or even an impediment to clear theorizing. I explore various reasons that reversion might have (...) appeared to be a central problem for this first group of figures, focusing on their commitment to a variety of conceptual features of evolutionary theory; discuss why reversion might have then ceased to be an interesting phenomenon; and, finally, close with some more general thoughts about the death of scientific problems. (shrink)

One claim found in the received historiography of the biometrical school (comprised primarily of Francis Galton, Karl Pearson, and W. F. R. Weldon) is that one of the biometricians' great flaws was their inability to look past their population-focused, statistical, gradualist understanding of evolutionary change – which led, in part, to their ignoring developments in cellular biology around 1900. I will argue, on the contrary, that the work of the biometricians was, from its earliest days, fundamentally concerned with connections between (...) statistical patterns of inheritance and the underlying cellular features that gave rise to them. Such work remained current with contemporary knowledge of chromosomes, cytology, and development; in this article, I explore the first case. The biometricians were thus well positioned to understand the relationship between the patterns of Mendelian inheritance and the statistical distributions with which they primarily occupied themselves. Ignorance of this connection, then, is not the reason why they rejected Mendelism. Further, both Galton and Weldon – though each in their own unique way – decided to turn to biological detail as a way to better justify the generality of their statistical approaches to heredity. Perhaps paradoxically, then, for these biometricians, detail offered an approach to theoretical generality. (shrink)

An important historical relation that has hardly been addressed is the influence of Prosper Lucas's Treatise on Natural Inheritance on the development of Charles Darwin's concepts related to inheritance. In this article we trace this historical connection. Darwin read Lucas's Treatise in 1856. His reading coincided with many changes concerning his prior ideas on the transmission and expression of characters. We consider that this reading led him to propose a group of principles regarding prepotency, hereditary diseases, morbid tendencies and atavism; (...) following Lucas, he called these principles: laws of inheritance. (shrink)

The cell is not only the structural, physiological, and developmental unit of life, but also the reproductive one. So far, however, this aspect of the cell has received little attention from historians and philosophers of biology. I will argue that cell theory had far-reaching consequences for how biologists conceptualized the reproductive relationships between germs and adult organisms. Cell theory, as formulated by Theodor Schwann in 1839, implied that this relationship was a specific and lawful one, that is, that germs of (...) a certain kind, all else being equal, would produce adult organisms of the same kind, and vice versa. Questions of preformation and epigenesis took on a new meaning under this presupposition. The question then became one of whether cells could be considered as autonomous agents producing adult organisms of a given species, or whether they were the product of external, organizing forces and thus only a stage in the development of the whole organism. This question became an important issue for nineteenth-century biology. As I will demonstrate, it was the view of cells as autonomous agents which helped both Charles Darwin and Gregor Mendel to think of inheritance as a lawful process. (shrink)

The cell is not only the structural, physiological, and developmental unit of life, but also the reproductive one. So far, however, this aspect of the cell has received little attention from historians and philosophers of biology. I will argue that cell theory had far-reaching consequences for how biologists conceptualized the reproductive relationships between germs and adult organisms. Cell theory, as formulated by Theodor Schwann in 1839, implied that this relationship was a specific and lawful one, that is, that germs of (...) a certain kind, all else being equal, would produce adult organisms of the same kind, and vice versa. Questions of preformation and epigenesis took on a new meaning under this presupposition. The question then became one of whether cells could be considered as autonomous agents producing adult organisms of a given species, or whether they were the product of external, organizing forces and thus only a stage in the development of the whole organism. This question became an important issue for nineteenth-century biology. As I will demonstrate, it was the view of cells as autonomous agents which helped both Charles Darwin and Gregor Mendel to think of inheritance as a lawful process. (shrink)

In the period 1875-1920, a debate about the generality and applicability of evolutionary theory to all organisms was motivated by work on unicellular ciliates like Paramecium because of their peculiar nuclear dualism and life cycles. The French cytologist Emile Maupas and the German zoologist August Weismann argued in the 1880s about the evolutionary origins and functions of sex (which in the ciliates is not linked to reproduction), and death (which appeared to be the inevitable fate of lineages denied sexual conjugation), (...) an argument rooted in the question of whether the ciliates and their processes where homologous to other cellular organisms. In the beginning of the twentieth century, this question of homology came to be less important as the ciliates were used by the British protozoologist Clifford Dobell and the American zoologist Herbert Spencer Jennings to study evolutionary processes in general rather than problems of development and cytology. For them, homology mattered less than analogy. This story illustrates two partially distinct problems in evolutionary biology: first, the question of whether all living things have common features and origins; and second, whether their history and current nature can be described by identical mechanisms. Where Maupas (contra Weismann) made the ciliates qualitatively the same as all other organisms in order to create a cohesive evolutionary theory for biology, Jennings and Dobell made them qualitatively different in order to achieve the same end. (shrink)

This paper argues that our modern concept of biological heredity was first clearly introduced in a theoretical and practical setting by the generation of French physicians that were active between 1810 and 1830. It describes how from a traditional focus on hereditary transmission of disease, influential French medical men like Esquirol, Fodéré, Piorry, Lévy, moved towards considering heredity a central concept for the conception of the human bodily frame, and its set of physical and moral dispositions. The notion of heredity (...) as a natural force, with a wide ranging capabilities of transmitting differentially both fundamental and accidental characters was generalized by that generation of physicians with the help of contemporary naturalists and physiologists. By 1830 the term hérédité was widespread, and it shared the explanatory and semantic qualities of traditional medical concepts like constitution and temperament. An analysis is given of the main developments that led to the conception of biological (including human) bodies as consisting of a layered, hierarchical organization of characters, differentially affected by the laws of conservation (Heredity) and change (Inneity, Variation). The mid-century work of the French physician Prosper Lucas, Traité Philosophique et Physiologique de L'Hérédité Naturelle, is shown to be the culmination of the efforts of several generations of French physicians towards having a feasible, complexly structured notion of how heredity works. (shrink)

Textbook descriptions of the foundations of Genetics give the impression that besides Mendel’s no other research on heredity took place during the nineteenth century. However, the publication of the Origin of Species in 1859, and the criticism that it received, placed the study of heredity at the centre of biological thought. Consequently, Herbert Spencer, Charles Darwin himself, Francis Galton, William Keith Brooks, Carl von Nägeli, August Weismann, and Hugo de Vries attempted to develop theories of heredity under an evolutionary perspective, (...) and they were all influenced by each other in various ways. Nonetheless, only Nägeli became aware of Mendel’s experimental work; it has also been questioned whether Mendel even had the intention to develop a theory of heredity. In this article, a short presentation of these theories is made, based on the original writings. The major aim of this article is to suggest that Mendel was definitely not the only one studying heredity before 1900, if he even did this, as may be inferred by textbooks. Although his work had a major impact after 1900, it had no impact during the latter half of the nineteenth century when an active community of students of heredity emerged. Thus, textbooks should not only present the work of Mendel, but also provide a wider view of the actual history and a depiction of science as a social process. (shrink)

Although it has not been much noticed, reproduction is the central theme of François Jacob’s important history of biology, La logique du vivant (The Logic of Life). In a book ostensibly devoted to heredity, this molecular biologist had reproduction integrate levels of organization from organisms to molecules and play a major role in each historical transition between them, not just in the influential argument for a shift “from generation to reproduction.” Moreover, I claim, La logique was the first general history (...) of (research on) reproduction. Earlier histories had tackled only aspects of generation and reproduction; none historicized the latter. Jacob’s innovation came out of the rising prominence of reproduction in neo-Darwinist studies of populations and in his own field, the cellular genetics of bacteria and their viruses, and engagement with history and philosophy of science. The discovery of bacterial “sex” enabled microbial genetics but, ironically, led Jacob to a sexless concept of reproduction. This approach molded a history that reached across the living world but slighted sex and the complexities of reproduction in multicellular organisms. Readers can learn from its strength and limitations. (shrink)

Bernard Norton's friends in the history of science have had many reasons for commemorating, with admiration and affection, not only his research and teaching but no less his conversation and his company. One of his most estimable traits was his refusal to beat about the bush in raising the questions he thought worthwhile pursuing. I still remember discoursing at Pittsburgh on Darwin's route to his theory of natural selection, and being asked at the end by Bernard what were Darwin's views (...) on heredity. I answered with the conventional waffle to the effect that the theory concerned the populational fate rather than the individual production and transmission of heritable variation, so that whatever views Darwin had on heredity had only a subsidiary place in his theorizing. Bernard was not fooled. ‘I would have thought’, he said, ‘that in order to understand anyone's theorising about evolution it would be necessary to look at his views on heredity’. (shrink)

This article considers the reception of British cytogeneticist C.D. Darlington's controversial 1932 book, Recent Advances in Cytology. Darlington's cytogenetic work, and the manner in which he made it relevant to evolutionary biology, marked an abrupt shift in the status and role of cytology in the life sciences. By focusing on Darlington's scientific method -- a stark departure from anti-theoretical, empirical reasoning to a theoretical and speculative approach based on deduction from genetic first principles -- the article characterises the relationships defining (...) the "disciplinary landscape" of the life sciences of the time, namely those between cytology, genetics, and evolutionary theory. (shrink)

This article considers the reception of British cytogeneticist C.D. Darlington's controversial 1932 book, Recent Advances in Cytology. Darlington's cytogenetic work, and the manner in which he made it relevant to evolutionary biology, marked an abrupt shift in the status and role of cytology in the life sciences. By focusing on Darlington's scientific method -- a stark departure from anti-theoretical, empirical reasoning to a theoretical and speculative approach based on deduction from genetic first principles -- the article characterises the relationships defining (...) the "disciplinary landscape" of the life sciences of the time, namely those between cytology, genetics, and evolutionary theory. (shrink)

Stem cells did not become a proper research object until the 1960 s. Yet the term and the basic mind-set—namely the conception of single undifferentiated cells, be they embryonic or adult, as the basic units responsible for a directed process of development, differentiation and increasing specialisation—were already in place at the end of the nineteenth century and then transmitted on a non-linear path in the form of tropes and diagrams. Ernst Haeckel and August Weismann played a special role in this (...) story. The first coined the term Stammzelle (stem cell), the second was the author of the first cellular stem-tree diagram. Even today, I shall argue, the understanding of stem cells, especially the popular perception, is to a large extent a Haeckelian–Weismannian one. After having demonstrated this, by analysing the terminology, in this essay I will focus on the use of cytogenetic tree diagrams between 1892 and 1925 and on the tacit understanding of stem cells that they transmit. (shrink)