Biological atomism postulates that all life is composed of elementary and indivisible vital units. The activity of a living organism is thus conceived as the result of the activities and interactions of its elementary constituents, each of which individually already exhibits all the attributes proper to life. This paper surveys some of the key episodes in the history of biological atomism, and situates cell theory within this tradition. The atomistic foundations of cell theory are subsequently dissected and discussed, together with (...) the theory’s conceptual development and eventual consolidation. This paper then examines the major criticisms that have been waged against cell theory, and argues that these too can be interpreted through the prism of biological atomism as attempts to relocate the true biological atom away from the cell to a level of organization above or below it. Overall, biological atomism provides a useful perspective through which to examine the history and philosophy of cell theory, and it also opens up a new way of thinking about the epistemic decomposition of living organisms that significantly departs from the physicochemical reductionism of mechanistic biology. (shrink)

Since the discovery of the double helical structure of DNA, the standard account of the inheritance of features has been in terms of DNA-copying and DNA-transmission. This theory is just a version of the old theory according to which the inheritance of features is explained by the transfer at conception of some developmentally privileged material from parents to offspring. This paper does the following things: (1) it explains what the inheritance of features is; (2) it explains how the DNA-centric theory (...) emerged; (3) it clarifies the relation between the DNA-centric theory and the ‘unfolding’ theory of development; (4) it argues that (given what we now know about developmental processes and genetic activity) the DNA-centric theory should be abandoned in favour of a pluralistic (but not holistic) theory of the inheritance of features. According to this pluralistic theory, the reliable reoccurrence of phenotypes must be explained by appealing not only to processes responsible for the reliable reoccurrence of genetic developmental factors but also to processes responsible for the reliable reoccurrence (or persistence) of nongenetic developmental factors. (shrink)

According to a classical narrative, early geneticists, failing to see how Mendelism provides the missing pieces of Darwin’s theory, rejected gradual changes and advocated an implausible yet briefly popular view of evolution-by-mutation; after decades of delay (in which synthesis was prevented by personal conflicts, disciplinary rivalries, and anti-Darwinian animus), Darwinism emerged on a new Mendelian basis. Based on the works of four influential early geneticists – Bateson, de Vries, Morgan and Punnett –, and drawing on recent scholarship, we offer an (...) alternative that turns the classical view on its head. For early geneticists, embracing discrete inheritance and the mutation theory (for the origin of hereditary variation) did not entail rejection of selection, but rejection of Darwin’s non-Mendelian views of heredity and variation, his doctrine of natura non facit saltum, and his conception of “natural selection” as a creative force that shapes features out of masses of infinitesimal differences. We find no evidence of a delay in synthesizing mutation, rules of discrete inheritance, and selection in a Mendelian-Mutationist Synthesis. Instead, before 1918, early geneticists had conceptualized allelic selection, the Hardy–Weinberg equilibrium, the evolution of a quantitative trait under selection, the probability of fixation of a new mutation, and other key innovations. Contemporary evolutionary thinking seems closer to their more ecumenical view than to the restrictive mid-twentieth-century consensus known as the Modern Synthesis. (shrink)

The hologenome concept of evolution is a hypothesis about the evolution of animals and plants. It asserts that the evolution of animals and plants was partially triggered by their interactions with their symbiotic microbiomes. In that vein, the hologenome concept posits that the holobiont (animal host + symbionts of the microbiome) is a unit of selection. -/- The hologenome concept has been severely criticized on the basis that selection on holobionts would only be possible if there were a tight transgenerational (...) host-genotype-to-symbiont-genotype connection. As our current evidence suggests that this is not the case for most of the symbiont species that compose the microbiome of animals and plants, the opportunity for holobiont selection is very low in relation to the opportunity for selection on each of the species that compose the host microbiome. Therefore, holobiont selection will always be disrupted ‘from below’, by selection on each of the species that compose the microbiome. -/- This thesis constitutes a conceptual effort to defend philosophically the hologenome concept. I argue that the criticism according to which holobiont selection requires tight transgenerational host-genotype-to-symbiont-genotype connection is grounded on a metaphysical view of the world according to which the biological hierarchy needs to be nested, such that each new level of selection includes every entity from below. Applied to hologenomes, it entails that the hologenome is a collection of genomes, and selection of hologenomes is assumed to entail cospeciation of the host with the species that constitute its microbiome. -/- Against that interpretation, I propose the ‘stability of traits’ account, according to which hologenome evolution is the result of the action of natural selection in a non-nested hierarchical world. In that vein, hologenome evolution does not entail cospeciation, and thus it does not require tight transgenerational host-genotype-to-symbiont-genotype connection. By embracing a multilevel selection perspective, I argue that hologenome evolution results from the simultaneous action of natural selection on each of the lineages that compose the microbiome, and on the assemblage composed by the host genome plus the functional traits of its microbiome. Hologenome selection occurs when the evolution of the traits of the microbiome result from their effects on the fitness of the host, and it can take the form of multilevel selection 1, or multilevel selection 2. In both cases, hologenome selection entails the evolution of microbiome traits, as well as evolution of the host genome, rather than cospeciation of lineages. (shrink)

Although the theory of memetics appeared highly promising at the beginning, it is no longer considered a scientific theory among contemporary evolutionary scholars. This study aims to compare the genealogy of memetics with the historically more successful gene-culture coevolution theory. This comparison is made in order to determine the constraints that emerged during the internal development of the memetics theory that could bias memeticists to work on the ontology of meme units as opposed to hypotheses testing, which was adopted by (...) the gene-culture scholars. I trace this problem back to the diachronic development of memetics to its origin in the gene-centered anti-group-selectionist argument of George C. Williams and Richard Dawkins. The strict adoption of this argument predisposed memeticists with the a priori idea that there is no evolution without discrete units of selection, which in turn, made them dependent on the principal separation of biological and memetic fitness. This separation thus prevented memeticists from accepting an adaptationist view of culture which, on the contrary, allowed gene-culture theorists to attract more scientists to test the hypotheses, creating the historical success of the gene-culture coevolution theory. (shrink)

This paper describes the historical background and early formation of Wilhelm Johannsen's distinction between genotype and phenotype. It is argued that contrary to a widely accepted interpretation his concepts referred primarily to properties of individual organisms and not to statistical averages. Johannsen's concept of genotype was derived from the idea of species in the tradition of biological systematics from Linnaeus to de Vries: An individual belonged to a group - species, subspecies, elementary species - by representing a certain underlying type. (...) Johannsen sharpened this idea theoretically in the light of recent biological discoveries, not least those of cytology. He tested and confirmed it experimentally combining the methods of biometry, as developed by Francis Galton, with the individual selection method and pedigree analysis, as developed for instance by Louis Vilmorin. The term "genotype" was introduced in W. Johannsen's 1909 treatise, but the idea of a stable underlying biological "type" distinct from observable properties was the core idea of his classical bean selection experiment published 6 years earlier. The individual ontological foundation of population analysis was a self-evident presupposition in Johannsen's studies of heredity in populations from their start in the early 1890s till his death in 1927. The claim that there was a "substantial but cautious modification of Johannsen's phenotype-genotype distinction" from a statistical to an individual ontological perspective derives from a misreading of the 1903 and 1909 texts. The immediate purpose of this paper is to correct this reading of the 1903 monograph by showing how its problems and results grow out of Johannsen's earlier work in heredity and plant breeding. Johannsen presented his famous selection experiment as the culmination of a line of criticism of orthodox Darwinism by William Bateson, Hugo de Vries, and others. They had argued that evolution is based on stepwise rather than continuous change in heredity. Johannsen's paradigmatic experiment showed how stepwise variation in heredity could be operationally distinguished from the observable, continuous morphological variation. To test Galton's law of partial regression, Johannsen deliberately chose pure lines of self-fertilizing plants, a pure line being the descendants in successive generations of one single individual. Such a population could be assumed to be highly homogeneous with respect to hereditary type, and Johannsen found that selection produced no change in this type. Galton, he explained, had experimented with populations composed of a number of stable hereditary types. The partial regression which Galton found was simply an effect of selection between types, increasing the proportion of some types at the expense of others. (shrink)

The paper aims at proposing an extended notion of epigenesis acknowledging an actual causal import to the phenotypic dimension for the evolutionary diversification of life forms. “Introductory remarks” section offers introductory remarks on the issue of epigenesis contrasting it with ancient and modern preformationist views. In “Transmutation of forms: phenotypic variation, diversification, and complexification” section we propose to intend epigenesis as a process of phenotypic formation and diversification dependent on environmental influences, independent of changes in the genomic nucleotide sequence, and (...) occurring during the whole life span. Then, “Evolvability and phenotypic evolution” section focuses on phenotypic plasticity and offers an overview of basic properties that allows biological systems to be evolvable—i.e. to have the potentiality of producing phenotypic variation. Successively, the emphasis is put on environmentally-induced modifications in the regulation of gene expression giving rise to phenotypic variation and diversification. After some brief considerations on the debated issue of epigenetic inheritance, the issue of culture is considered. The key point is that, in the case of humans and of the evolutionary history of the genus Homo at least, the environment is also, importantly, the cultural environment. Thus, “Bio-cultural feedback” section argues that a bio-cultural feedback should be acknowledged in the “epigenic” processes leading to phenotypic diversification and innovation in Homo evolution. Finally, “Brain plasticity and cultural neural reuse” section introduces the notion of “cultural neural reuse”, which refers to phenotypic/neural modifications induced by specific features of the cultural environment that are effective in human cultural evolution without involving genetic changes. Therefore, cultural neural reuse may be regarded as a key instance of the bio-cultural feedback and ultimately of the extended notion of epigenesis proposed in this work. (shrink)

The new discipline of exobiology formed from the intertwining of origin of life research with the search for life or its building blocks on other planets, from 1957-1973. The field was inherently highly interdisciplinary, yet it coalesced very quickly and was responsible in its first twenty years for numerous important contributions to twentieth century life science and planetary sciences such as climatology, the study of mass extinctions, etc. NASA played a very important role in catalyzing the rapid consolidation of exobiology, (...) both through research grants and through sponsored meetings that overcame disciplinary boundaries, bringing together scientists from diverse backgrounds. The presence of a handful of prominent senior scientists such as Joshua Lederberg, Melvin Calvin and Norman Horowitz helped gain credibility for exobiology, in the face of criticism and competition from existing life sciences disciplines. Tensions within the exobiology research community and between NASA-funded science and the academic research community are explored, as are such milestones of discipline formation as journals and professional societies. (shrink)

John Richardson (2002, 2004) argues that Nietzsche’s use of teleological notions, such as the “will to power” and psychological “drives,” can be naturalized within the Darwinian framework of natural selection. Although this ambitious project has merit, the Darwinian framework does not provide the strong teleology necessary to interpret Nietzsche’s explanatory project. Examining the logic of selection, the conceptual limitations on biological functions, and the evidential demands that must be met to deploy evolutionary theory show that Nietzsche’s explanatory project does not (...) cohere with the Darwinian framework. Thus, coherence with currently accepted evolutionary theory should not constrain the philosophical project of interpretation in this case. (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)

Pluralism is popular among philosophers of biology. This essay argues that negative judgments about universal biology, while understandable, are very premature. Familiar life on Earth represents a single example of life and, most importantly, there are empirical as well as theoretical reasons for suspecting that it may be unrepresentative. Scientifically compelling generalizations about the unity of life must await the discovery of forms of life descended from an alternative origin, the most promising candidate being the discovery of extraterrestrial life. Nonetheless, (...) in the absence of additional examples of life, we are best off exploring the microbial world for promising explanatory concepts, principles, and mechanisms rather than prematurely giving up on universal biology. Unicellular microbes are by far the oldest, metabolically most diverse, and environmentally tolerant form of life on our planet. Yet somewhat ironically, much of our theorizing about life still implicitly privileges complex multicellular eukaryotes, which are now understood to be highly specialized, fragile latecomers to Earth. The problem with pursuing a pluralist approach to understanding life is that it is likely to blind us to the significance of just those entities and causal processes most likely to shed light on the underlying nature of life. (shrink)

Philosophers of biology, along with everyone else, generally perceive life to fall into two broad categories, the microbes and macrobes, and then pay most of their attention to the latter. ‘Macrobe’ is the word we propose for larger life forms, and we use it as part of an argument for microbial equality. We suggest that taking more notice of microbes – the dominant life form on the planet, both now and throughout evolutionary history – will transform some of the philosophy (...) of biology’s standard ideas on ontology, evolution, taxonomy and biodiversity. We set out a number of recent developments in microbiology – including biofilm formation, chemotaxis, quorum sensing and gene transfer – that highlight microbial capacities for cooperation and communication and break down conventional thinking that microbes are solely or primarily single-celled organisms. These insights also bring new perspectives to the levels of selection debate, as well as to discussions of the evolution and nature of multicellularity, and to neo-Darwinian understandings of evolutionary mechanisms. We show how these revisions lead to further complications for microbial classification and the philosophies of systematics and biodiversity. Incorporating microbial insights into the philosophy of biology will challenge many of its assumptions, but also give greater scope and depth to its investigations. (shrink)

"Greed, for lack of a better word, is good. Greed is right. Greed works. Greed clarifies, cuts through, and captures the essence of the evolutionary spirit." "From the outset [in 1976] the reviews were gratifyingly favorable and it [The Selfish Gene] was not seen, initially, as a controversial book. Its reputation for contentiousness took years to grow until, by now, it is widely regarded as a work of radical extremism. But over the very same years as the book’s reputation for (...) extremism has escalated, its actual content has seemed less and less extreme, more and more the common currency." How could it be that Dawkins’ most famous book became controversial, but not as a result of what it .. (shrink)

Historians and biologists identify the debate between mechanists and vitalists over the nature of life itself with the arguments of Driesch, Loeb, and other prominent voices. But what if the conversation was broader and the consequences deeper for the field? Following the suspicions of Joseph Needham in the 1930s and Francis Crick in the 1960s, we deployed tools of the digital humanities to an old problem in the history of biology. We analyzed over 31,000 peer-reviewed scientific papers and learned that (...) bioexceptionalism participated in a robust discursive landscape throughout subfields of the life sciences, occupied even by otherwise unknown biologists. (shrink)

In “The Origin of Species,” Darwin describes a hypothetical example illustrating that large, slowly reproducing mammals such as the elephant can reach very large numbers if population growth is not affected by regulating factors. The elephant example has since been cited in various forms in a wide variety of books, ranging from educational material to encyclopedias. However, Darwin’s text was changed over the six editions of the book, although some errors in the mathematics persisted throughout. In addition, full details of (...) the problem remained hidden in his correspondence with readers of the Origin. As a result, Darwin’s example is very often misinterpreted, misunderstood or presented as if it were a fact. We show that the population growth of Darwin’s elephant population can be modeled by the Leslie matrix method, which we generalize here to males as well. Darwin’s most often cited figure, about 19 million elephants after 750 years is not a typical outcome, actually a very unlikely result under more realistic, although still hypothetical situations. We provide a recursion formula suggesting that Darwin’s original model corresponds to a tribonacci series, a proof showing that sex ratio is constant over all age classes, and a derivation of a generating function of the sequence. (shrink)

Fundamental questions in evolution concern deep divisions in the living world and vertical versus horizontal information transfer. Two contrasting views are: (i) three superkingdoms Archaea, Eubacteria, and Eukarya based on vertical inheritance of genes encoding ribosomes; versus (ii) a prokaryotic/eukaryotic dichotomy with unconstrained horizontal gene transfer (HGT) among prokaryotes. Vertical inheritance implies continuity of cytoplasmic and structural information whereas HGT transfers only DNA. By hypothesis, HGT of the translation machinery is constrained by interaction between new ribosomal gene products and vertically (...) inherited cytoplasmic structure made largely of preexisting ribosomes. Ribosomes differentially enhance the assembly of new ribosomes made from closely related genes and inhibit the assembly of products from more distal genes. This hypothesis suggests experiments for synthetic biology: the ability of synthetic genomes to “boot,” i.e., establish hereditary continuity, will be constrained by the phylogenetic closeness of the cell “body” into which genomes are placed. (shrink)

RESUMENEl desarrollo embriológico es un fenómeno que ha inspirado la especulación filosófica desde temprano en la historia del pensamiento. Desde los tiempos de Aristóteles dos modelos conceptuales antitéticos se han utilizado tradicionalmente para comprender la embriogénesis: o el embrión posee ya una forma o estructura, o ésta se forma de nuevo en cada generación. Nuestro objetivo en este artículo es mostrar que el contraste entre la posición preformacionista y epigenética persiste a pesar de los formidables avances teóricos y experimentales de (...) la biología del desarrollo contemporánea. El preformacionismo y la epigénesis han perfeccionado constantemente sus posiciones en el curso de la historia con el fin de responder a los retos conceptuales de la época. Este continuo proceso de transformación ha dado lugar a una convergencia parcial entre las dos posiciones. Sin embargo, vamos a argumentar que, a pesar de los esfuerzos por conciliar ambas posturas, esta antinomia, que se erige como una de las más fundamentales de la biología, no será fácil de superar. ABSTRACTThe process of development of the embryo has inspired philosophical speculation since the advent of Western thought. Since the time of Aristotle two antithetical conceptual models have traditionally been used to understand embryogenesis: either form or structure is preformed in the embryo or it is newly formed with each generation. Our aim in this article is to show that the contrast between the pre-formationist and epigenetic positions persists despite the formidable theoretical and experimental advances of contemporary developmental biology. Pre-formationism and epigenesis have constantly refined their positions in the course of history in order to answer the conceptual challenges of each epoch. This process of continuous transformation has resulted in a partial convergence of the two positions. However, we shall argue that, despite the partial successes of this process of continuous convergence, one of the most fundamental antinomies in biology persists. (shrink)

It is of philosophical and epistemological interest to examine how Darwin conceived the process of division of labour within Natural History. Darwin observed the advantages brought by division of labour to the human economy, and considered that the principle of divergence within nature, which is, according to him, one of the two ‘keystones’ of his theory, gave comparable advantages. This led him to re-examine Milne-Edwards’ view on the notion of division of physiological labour, and to introduce this with modifications into (...) his naturalist writings. After a short review of the Darwinian historiography dealing with this issue, I first show the conceptual confusion into which Darwin plunges, when using a so-called economic argument to defend his thesis of the maximization of beings in a given territory due to division of labour. Following this I propose several hypotheses to explain these shifts, recurring in Darwin’s texts, from one conception and from one application to another, of the division of labour. (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)

After Darwin, experimental biology sought to unravel organisms. By the early twentieth century, organisms were broadly conceived as the product of their heredity and their environment. Much historical work has explored the scientific attack on the genotype, particularly through the new science of genetics. This article explores the tandem efforts to assert experimental control over the environment in which plants grew and developed. The case described here concerns the creation of the first phytotron at Caltech by botanist and plant physiologist (...) Frits Went. Opening in 1949, the phytotron was a plant laboratory that, across a series of rooms and chambers, kept genes constant while regulating and maintaining defined ranges of known environments. This article details the context in which the phytotron emerged, how the phytotron gained its sobriquet, and how it served to cement the “environment” as a category of biological knowledge. Describing the institutional context of Caltech, its interdisciplinary culture, and its encouragement of adopting technology into biological science, I argue that the phytotron and the commensurate category of the “environment” were the product of the familiar movement to integrate the physical and biological sciences. In addition, however, the creation of the phytotron was also a broader story of plant physiologists establishing a definition of the “environment” in both physical and technological terms. (shrink)

This paper examines the history of Japanese genetics in the 1920s to 1950s as seen through the work of Hitoshi Kihara, a prominent wheat geneticist as well as a leader in the development of the discipline in Japan. As Kihara's career illustrates, Japanese genetics developed quickly in the early twentieth century through interactions with biologists outside Japan. The interactions, however, ceased due to the war in the late 1930s, and Japanese geneticists were mostly isolated from outside information until the late (...) 1940s. During the isolation in wartime and under the postwar U.S. Occupation, Kihara adapted to political changes. During wartime, he developed a research institute focusing on applied biology of various crops, which conformed to the national need to address food scarcity. After the war, he led the campaign for the establishment of a national institute of genetics and negotiated with American Occupation officers. The Americans viewed this Japanese effort with suspicion because of the rising popularity of the controversial theory of the Russian agronomist, Trofim Lysenko, in Japan. The institute was approved in 1949 partly because Kihara was able to bridge the gap between the American and Japanese sides. With Kihara's flexible and generous leadership, Japanese genetics steadily developed, survived the wartime, and recovered quickly in the postwar period. The article discusses Kihara's interest in cytoplasmic inheritance and his synthetic approach to genetics in this political context, and draws attention to the relation between Kihara's genetics and agricultural practice in Japan. (shrink)

In 1877, the young Alfred Espinas defended a philosophical study, ‘doctorat ès lettres’, at the Sorbonne University, entitled Des Sociétés animales. This was to become one of the principal sources of French organicist sociology. The paradox, however, is that this work seems to be fundamentally a study of natural science. Espinas tried to justify his position theoretically through two types of reciprocally exclusive and uncomplementary arguments. The first one consists in showing that only certain kinds of animal groupings belong legitimately, (...) if not academically, to sociology. The second one has a greater audacious and uncontrollable dimension. It consists in asserting that all pluricellular organisms are true societies. In this article we first focus on the role played by cellular theory in such a ‘sociologization’ of biology, concerning the extension of sociology. Second, we examine the importance of such confusion within the fields of extension, in aiming to explain the conceptual transfers between social sciences and life sciences, in the second half of the 19th century. (shrink)