Richard Woltereck was a German zoologist and hydrobiologist who studied aquatic animals and extended the concept of Reaktionsnorm (norm of reaction) to the study of genetics. He also provided some of the first experimental evidence for the early twentieth-century embryological theory of heredity known as cytoplasmic inheritance. Through experiments on the water flea, Daphnia, Woltereck investigated whether variation produced by environmental impacts on development could play a role in heredity and evolution. Woltereck's research emphasized the importance of environment and development in Wilhelm Johannsen's concepts of genotype and phenotype. Biologists throughout the twentieth century used Woltereck's concept of Reaktionsnorm to develop theories and experiments to explain the evolution of adaptive developmental responses to environmental conditions. Later in his career, Woltereck developed a theory of heredity that sought to reconcile embryological concepts, such as regulation and body plans, with Mendelian heredity and Darwinian evolution by natural selection.
Richard Woltereck first described the concept of Reaktionsnorm (norm of reaction) in his 1909 paper 'Weitere experimentelle Untersuchungen uber Art-veranderung, speziell uber das Wesen quantitativer Artunterschiede bei Daphniden' ('Further investigations of type variation, specifically concerning the nature of quantitative differences between varieties of Daphnia'). This concept refers to the ways in which the environment can alter the development of an organism, and its adult characteristics. Woltereck conceived of the Reaktionsnorm as the full range of potentialities latent in a single genotype, evocable by the environmental circumstances of a developing organism. Biologists used variants of Woltereck's concept of Reaktionsnorm, often called the reaction norm or norm of reaction, throughout the twentieth century in attempts to explain how developmental responses to the environment can evolve, and even alter the tempo and direction of evolutionary change.
In 2008 researchers Daniel Warner and Richard Shine tested the Charnov-Bull model by conducting experiments on the Jacky dragon (Amphibolurus muricatus), in Australia. Their results showed that temperature-dependent sex determination(TSD) evolved in this species as an adaptation to fluctuating environmental temperatures. The Charnov-Bull model, proposed by Eric Charnov and James Bull in 1977, described the evolution of TSD, although the model was, for many years, untested. Many reptiles and some fish exhibit non-genetic sex determination, in which an embryos' environment can influence the sex of the adult organism. Environmental conditions such as humidity or population density can alter sex in some organisms, and a widespread form of non-genetic sex determination is temperature-dependent sex determination. TSD reveals how embryonic development can contribute to the evolution of physiological processes. Researchers have documented TSD in a wide range of species, and they continue to investigate how such a sex determining system has evolved.
Ernst Heinrich Philipp August Haeckel was a prominent comparative anatomist and active lecturer in the late nineteenth and early twentieth centuries. He is most well known for his descriptions of phylogenetic trees, studies of radiolarians, and illustrations of vertebrate embryos to support his biogenetic law and Darwin's work with evolution. Haeckel aggressively argued that the development of an embryo repeats or recapitulates the progressive stages of lower life forms and that by studying embryonic development one could thus study the evolutionary history of life on earth.
Mutual Affinities of Organic Beings: Morphology: Embryology: Rudimentary Organs is the thirteenth chapter of Charles Darwin's book The Origin of Species, first published in England in 1859. The book details part of Darwin's argument for the common ancestry of life and natural selection as the cause of speciation. In this chapter, Darwin summarizes the evidence for evolution by connecting observations of development in organisms to the processes of natural selection. Darwin shows how the theory of special creation, which claims that God directly created all organisms in their current form, is inferior to the theory of natural selection for its ability to explain the diversity of life. In this chapter, Darwin also discusses classification and homology as they relate to natural selection.
Ann Campbell Burke examines the development and evolution of vertebrates, in particular, turtles. Her Harvard University experiments, described in Development of the Turtle Carapace: Implications for the Evolution of a Novel Bauplan, were published in 1989. Burke used molecular techniques to investigate the developmental mechanisms responsible for the formation of the turtle shell. Burke's work with turtle embryos has provided empirical evidence for the hypothesis that the evolutionary origins of turtle morphology depend on changes in the embryonic and developmental mechanisms underpinning shell production.
Edward Drinker Cope studied fossils and anatomy in the US in the late nineteenth century. Based on his observations of skeletal morphology, Cope developed a novel mechanism to explain the law of parallelism, the idea that developing organisms successively pass through stages resembling their ancestors. Others had proposed the addition of new body forms at the end of an individual organism's developed as a mechanism through which new species arose, but those proposals relied on changes in the lengths of gestation or incubation. Cope proposed that a change in the growth rate of an embryo or fetus would allow the formation of new body forms while gestation or incubation periods remained constant. Thus, the growth of an embryo or fetus must become faster or slower to alter the number of stages during growth. Many paleontologists and geologists of the time, including Henry Fairfield Osborn and Louis Agassiz, accepted Cope's mechanisms of evolution as alternatives to natural selection as the causes generating new species, yet Cope proposed his mechanism solely as a way by which new genera arise. He advocated the neo-Lamarckian theory that new species evolve through the inheritance of acquired characteristics.
In 1962 the journal Acta Biotheoretica published the final work of the biologist Edward Stuart Russell, a full eight years after his death. Entitled The Diversity of Animals: an Evolutionary Study, this short, unfinished manuscript on evolution received little recognition in the scientific presses despite both its technical discussion of adaptations in decapods (crabs, shrimp, etc.) and its different approach to evolutionary theory. The precise reason for this neglect is unclear. This book is a continuation of Russell's philosophical perspective, organicism, an interpretation that focuses on the organism as the primary unit of analysis for the biological sciences. Russell first argued for this position in several of his earlier works, such as The Interpretation of Development and Heredity (1930) and The Directiveness of Organic Activities (1946). What was new in The Diversity of Animals lies in Russell's orthogenetic theory of evolution. By "orthogenetic" he means evolutionary change in definite directions. The overall thesis of this work is that transformations in evolution that occur in early ontogenesis, or development, are the best explanation for most diversity in nature. The consequence of Russell's argument is that an understanding of development is fundamental to an explanation of the major transformations in the evolutionary history of life.
The Law of Acceleration of Growth is a theory proposed by Edward Drinker Cope in the US during the nineteenth century. Cope developed it in an attempt to explain the evolution of genera by appealing to changes in the developmental timelines of organisms. Cope proposed this law as an additional theory to natural selection. He argued that the evolution of genera, the more general groups within which biologists group species, occurs when the individual in a species move through developmental stages faster than did their ancestors, but within the same fixed period of gestation, and thus can undergo new developmental stages and develop new traits. The Law of Acceleration compliments Cope's Law of Retardation of Growth. He described the later law as the process by which organisms revert to an ancestral stage. In these cases, forces suppress the most recent traits or stages common to the development of individuals from different species within the same genus. Cope described evolution as progressive, following a predetermined path, a perspective about evolution sometimes called orthogenetic. Cope's was one among many orthogenic theories in the second half of the nineteenth century. Furthermore, the theory was part of a trend in nineteenth century in which some biologists claimed that the changes in developmental timing of organisms could explain large changes in biological forms throughout natural history.
Cellular automata (CA) are mathematical models used to simulate complex systems or processes. In several fields, including biology, physics, and chemistry, CA are employed to analyze phenomena such as the growth of plants, DNA evolution, and embryogenesis. In the 1940s John von Neumann formalized the idea of cellular automata in order to create a theoretical model for a self-reproducing machine. Von Neumann's work was motivated by his attempt to understand biological evolution and self-reproduction.