During the 1950s, the English biologist C. H. Waddington called for an extended synthesis based on his research on epigenetics and genetic assimilation. An extended synthesis was also proposed by the Austrian zoologist Rupert Riedl, with the study of evolvability. In 1978, Michael J. D. White wrote about an extension of the modern synthesis based on new research from speciation.
1980s: punctuated equilibrium
In the 1980s, the American palaeontologists Stephen Jay Gould and Niles Eldredge argued for an extended synthesis based on their idea of punctuated equilibrium, the role of species selection shaping large scale evolutionary patterns and natural selection working on multiple levels extending from genes to species. The ethologist John Endler wrote a paper in 1988 discussing processes of evolution that he felt had been neglected.
Contributions from evolutionary developmental biology
Some researchers in the field of evolutionary developmental biology proposed another synthesis. They argue that the modern and extended syntheses should mostly center on genes and suggest an integration of embryology with molecular genetics and evolution, aiming to understand how natural selection operates on gene regulation and deep homologies between organisms at the level of highly conserved genes, transcription factors and signalling pathways. By contrast, a different strand of evo-devo following an organismal approach contributes to the extended synthesis by emphasizing developmental bias, evolvability, and inherency of form as primary factors in the evolution of complex structures and phenotypic novelties. The principal focus of dispute with Neo-Darwinists is over the source of variation on which Natural Selection can operate. Supporters of the Extended Synthesis deny that these arise from "random copying errors" in DNA replication. In fact they often assert that development of an improved organism by such a mechanism - even occasionally - would violate Shannon's principles of Information Theory.
Recent history
The idea of an extended synthesis was relaunched in 2007 by Massimo Pigliucci, and Gerd B. Müller with a book in 2010 titled Evolution: The Extended Synthesis, which has served as a launching point for work on the extended synthesis. This includes:
The role of prior configurations, genomic structures, and other traits in the organism in generating evolutionary variations.
How increasing dimensionality of fitness landscapes affects our view of speciation.
The role of multilevel selection in the major evolutionary transitions.
How organisms modify the environments they belong to through niche construction.
Other processes such as evolvability, phenotypic plasticity, reticulate evolution, sex evolution and symbiogenesis are said by proponents to have been excluded or missed from the modern synthesis. The goal of Piglucci's and Müller's extended synthesis is to take evolution beyond the gene-centered approach of population genetics to consider more organism- and ecology-centered approaches. Many of these causes are currently considered secondary in evolutionary causation, and proponents of the extended synthesis want them to be considered first-class evolutionary causes. The biologist Eugene Koonin wrote in 2009 that "the new developments in evolutionary biology by no account should be viewed as refutation of Darwin. On the contrary, they are widening the trails that Darwin blazed 150 years ago and reveal the extraordinary fertility of his thinking."
Predictions
The extended synthesis is characterized by its additional set of predictions that differ from the standard modern synthesis theory:
changes in phenotype are predominantly positive, rather than neutral
changes in phenotype are induced in many organisms, rather than one organism
revolutionary change in phenotype can occur through mutation, facilitated variation or threshold events
repeated evolution in isolated populations can be by convergent evolution or developmental bias
adaptation can be caused by natural selection, environmental induction, non-genetic inheritance, learning and cultural transmission
rapid evolution can result from simultaneous induction, natural selection and developmental dynamics
biodiversity can be affected by features of developmental systems such as differences in evolvability
heritable variation is directed towards variants that are adaptive and integrated with phenotype
niche construction is biased towards environmental changes that suit the constructor's phenotype, or that of its descendants, and enhance their fitness
The extended evolutionary synthesis is currently being tested by a group of scientists from eight institutions in Britain, Sweden and the United States. The £7.7 million project is supported by a £5.7 million grant from the John Templeton Foundation. The project is headed by Kevin N. Laland at the University of St Andrews and Tobias Uller at Lund University. According to Laland what the extended synthesis "really boils down to is recognition that, in addition to selection, drift, mutation and other established evolutionary processes, other factors, particularly developmental influences, shape the evolutionary process in important ways."
Status
Biologists disagree on the need for an extended synthesis. Opponents contend that the modern synthesis is able to fully account for the newer observations, whereas others criticize that the Extended synthesis is not radical enough. Proponents think that the conceptions of evolution at the core of the modern synthesis are too narrow. Proponents argue that even when the modern synthesis allows for the ideas in the extended synthesis, using the modern synthesis affects the way that biologists think about evolution. For example, Denis Noble says that using terms and categories of the modern synthesis distort the picture of biology that modern experimentation has discovered. Proponents therefore claim that the extended synthesis is necessary to help expand the conceptions and framework of how evolution is considered throughout the biological disciplines.
