Antonine Nicoglou

This is an introduction to a collection of articles on the conceptual history of epigenesis, from Aristotle to Harvey, Cavendish, Kant and Erasmus Darwin, moving into nineteenth-century biology with Wolff, Blumenbach and His, and onto the twentieth century and current issues, with Waddington and epigenetics. The purpose of the topical collection is to emphasize how epigenesis marks the point of intersection of a theory of biological development and a theory of active matter. We also wish to show that the concept of epigenesis existed prior to biological theorization and that it continues to permeate thinking about development in recent biological debates.

In 1956, in his Principles of Embryology, Conrad Hal Waddington explained that the word “epigenetics” should be used to translate and update Wilhelm Roux’ German notion of “Entwicklungsmechanik” to qualify the studies focusing on the mechanisms of development. When Waddington mentioned it in 1956, the notion of epigenetics was not yet popular, as it would become from the 1980s. However, Waddington referred first to the notion in the late 1930s. While his late allusion clearly reveals that Waddington readily associated the notion of epigenetics with the developmental process, in the contemporary uses of the notion this developmental connotation seems to have disappeared. The advent and success of molecular biology have probably contributed to focusing biologists’ attention on the “genetic” or the “non-genetic” over the “developmental”. In the present paper, I first examine the links that exist, in Waddington’s work, between the classical notion of epigenesis in embryology and those of epigenetics that Waddington proposed to connect, and even synthesize, data both from embryology and genetics. Second, I show that Waddington’s own view of epigenetics has changed over time and I analyze how these changes appear through his many representations of the relationships between genetic signals and developmental processes.

The paper describes the context and the origin of a particular debate that concerns the evolution of phenotypic plasticity. In 1965, British biologist A. D. Bradshaw proposed a widely cited model intended to explain the evolution of norms of reaction, based on his studies of plant populations. Bradshaw’s model went beyond the notion of the “adaptive norm of reaction” discussed before him by Dobzhansky and Schmalhausen by suggesting that “plasticity” the ability of a phenotype to be modified by the environment should be genetically determined. To prove Bradshaw’s hypothesis, it became necessary for some authors to identify the pressures exerted by natural selection on phenotypic plasticity in particular traits, and thus to model its evolution. In this paper, I contrast two different views, based on quantitative genetic models, proposed in the mid-1980s: Russell Lande and Sara Via’s conception of phenotypic plasticity, which assumes that the evolution of plasticity is linked to the evolution of the plastic trait itself, and Samuel Scheiner and Richard Lyman’s view, which assumes that the evolution of plasticity is independent from the evolution of the trait. I show how the origin of this specific debate, and different assumptions about the evolution of phenotypic plasticity, depended on Bradshaw’s definition of plasticity and the context of quantitative genetics.

The concept of plasticity has always been present in the history of developmental biology, both within the theory of epigenesis and within morphogenesis studies. However this tradition relies also upon a genetic conception of plasticity. Founded upon the concepts of ‘‘phenotypic plasticity’’ and ‘‘reaction norm,’’ this genetic conception focuses on the array of possible phenotypic change in relation to diversified environments. Another concept of plasticity can be found in recent publications by some developmental biologists (Gilbert, West-Eberhard). I argue that these authors adopt a ‘‘broad conception of plasticity’’ that is closely related to a notion of development as something that is ongoing throughout an organism’s lifecycle, and has no clear-cut boundaries. However, I suggest that given a narrow conception of plasticity, one can define temporal boundaries for development that are linked to specific features of the morphological process, which are different from behavioral and physiological processes.

The biologist Mary Jane West-Eberhard publishes, in 2003, a book, entitled Developmental Plasticity and Evolution, in which a new synthetic approach, integrating development with evolution, is offered. For this reason, the book is seen as a piece of work in the field of Evolutionary developmental biology, commonly known as Evo Devo, whose aim is to synthesize data from both development and evolution. However, West-Eberhard's ambition is much higher in this book because she does not only focus on gathering the data from both fields, but she also wants to formalize a new synthetic theory of evolution, which includes development in its definition. She also explains that, in order to provide such a synthetic theory of evolution, she needs to offer an "inclusive definition of plasticity" in order to avoid "unnecessary distinctions at every turn" even if she assumes that such distinctions may be important for certain points. Our aim, in this article, is to explore the implications and consequences of West-Eberhard's "inclusive definition of plasticity" in such an attempt of a new synthesis of evolutionary theory, including development