At the turn of the twentieth century, William Bateson studied organismal variation and heredity of traits within the framework of evolutionary theory in England. Bateson applied Gregor Mendel's work to Charles Darwin's theory of evolution and coined the term genetics for a new biological discipline. By studying variation and advocating Mendelian genetics, Bateson furthered the field of genetics, encouraged the use of experimental methodology to study heredity, and contributed to later theories of genetic inheritance.

The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Programme, hereafter called The Spandrels, is an article written by Stephen J. Gould and Richard C. Lewontin published in the Proceedings of the Royal Society of London in 1979. The paper emphasizes issues with what the two authors call adaptationism or the adaptationist programme as a framework to explain how species and traits evolved. The paper is one in a series of works in which Gould emphasized the role of development in evolutionary theories. The article suggests that constraints on how organisms can develop and constraints on how species can evolve from others play a central role in explaining the how species and traits evolve. The authors note that organisms from different species develop as embryos through stages similar across species, genera, and higher classes. Gould and Lewontin hypothesize that those stages constrained the possible pathways of evolution and has therefore guided the history of life. Throughout the paper, the authors rely on analogy of some parts of organisms to architectural structures called spandrels, marked in this image as 'a'."

Paul Kammerer conducted experiments on amphibians and marine animals at the Vivarium, a research institute in Vienna, Austria, in the early twentieth century. Kammerer bred organisms in captivity, and he induced them to develop particular adaptations, which Kammerer claimed the organismss offspring would inherit. Kammerer argued that his results demonstrated the inheritance of acquired characteristics, or Lamarckian inheritance. The Lamarckian theory of inheritance posits that individuals transmit acquired traits to their offspring. Kammerer worked during a period in which scientists debated how variation between organisms and within species was caused, and how organisms could inherit that variation from their parents. Kammerer contended that the inheritance of acquired characteristics occurs during embryological development, but several scientists argued that he provided poor evidence for his claims.

In the early twentieth century, Paul Kammerer conducted a series of experiments to demonstrate that organisms could transmit characteristics acquired in their lifetimes to their offspring. In his 1809 publication, zoologist Jean-Baptiste Lamarck had hypothesized that living beings can inherit features their parents or ancestors acquired throughout life. By breeding salamanders, as well as frogs and other organisms, Kammerer tested Lamarck's hypothesis in an attempt to provide evidence for Lamarck's theory of the inheritance of acquired characteristics. In particular, Kammerer argued that the inheritance of acquired characteristics caused species to evolve, and he claimed that his results provided an explanation for evolutionary processes through developmental phenomena.

In the first decade of the twentieth century, Paul Kammerer, a zoologist working at the Vivarium in Vienna, Austria, conducted research on developmental mechanisms, including a series of breeding experiments on toads (Alytes obstetricans). Kammerer claimed that his results demonstrated that organisms could transmit acquired characteristics to their offspring. To explain how evolution occurred, biologist Jean-Baptiste Lamarck in France suggested in his 1809 book that offspring inherited the features their ancestors acquired throughout the lives of those ancestors, a process termed the inheritance of acquired characteristics. Kammerer conducted breeding experiments to test the theory of inheritance of acquired characteristics, which he said described the mechanics of evolution. Additionally, Kammerer's experiments aimed at explaining how development shaped evolutionary processes.

In the early twentieth century, Paul Kammerer, a zoologist working at the Vivarium in Vienna, Austria, experimented on sea-squirts (Ciona intestinalis). Kammerer claimed that results from his experiments demonstrated that organisms could transmit characteristics that they had acquired in their lifetimes to their offspring. Kammerer conducted breeding experiments on sea-squirts and other organisms at a time when Charles Darwin's 1859 theory of evolution lacked evidence to explain how offspring inherited traits from their parents. In 1809, zoologist Jean-Baptiste Lamarck in France theorized that living beings can inherit the features their parents or ancestors acquired during those ancestor's lifetime, a theory called the inheritance of acquired characteristics. Kammerer attempted to provide evidence for the theory of inheritance of acquired characteristics, which constituted, he argued, the mechanics of evolution. Kammerer claimed that his results could explain evolutionary processes through developmental phenomena.

Stephen Jay Gould studied snail fossils and worked at Harvard University in Cambridge, Massachusetts during the latter half of the twentieth century. He contributed to philosophical, historical, and scientific ideas in paleontology, evolutionary theory, and developmental biology. Gould, with Niles Eldredge, proposed the theory of punctuated equilibrium, a view of evolution by which species undergo long periods of stasis followed by rapid changes over relatively short periods instead of continually accumulating slow changes over millions of years. In his 1977 book, Ontogeny and Phylogeny, Gould reconstructed a history of developmental biology and stressed the importance of development to evolutionary biology. In a 1979 paper coauthored with Richard Lewontin, Gould and Lewonitn criticized many evolutionary bioligists for relying solely on adaptive evolution as an explanation for morphological change, and for failing to consider other explanations, such as developmental constraints.

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.

The sex of a reptile embryo partly results from the production of sex hormones during development, and one process to produce those hormones depends on the temperature of the embryo's environment. The production of sex hormones can result solely from genetics or from genetics in combination with the influence of environmental factors. In genotypic sex determination, also called genetic or chromosomal sex determination, an organism's genes determine which hormones are produced. Non-genetic sex determination occurs when the sex of an organism can be altered during a sensitive period of development due to external factors such as temperature, humidity, or social interactions. Temperature-dependent sex determination (TSD), where the temperature of the embryo's environment influences its sex development, is a widespread non-genetic process of sex determination among vertebrates, including reptiles. All crocodilians, most turtles, many fish, and some lizards exhibit TSD.