Extended Evolutionary Synthesis
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The extended evolutionary synthesis is an extension of the mid-20th century Modern Synthesis of evolutionary biology. It revisits the relative importance of different factors at play, revisits several assumptions of the original synthesis, and augments it with additional causative factors in evolution.
During the 1950s, biologist C. H. Waddington called for an extended synthesis based from his research on epigenetics and genetic assimilation. An extended synthesis was also proposed by 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.
In the 1980s, Stephen Jay Gould argued for an extended synthesis. This was based on his 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. Similar ideas were held by Niles Eldredge author of the book Unfinished Synthesis (1985).
Ethologist John Endler wrote a paper in 1988 discussing processes of evolution that he felt had been neglected. Others, such as researchers in the field of evolutionary developmental biology (evo-devo) called for a different extended synthesis based on the opinion that the modern synthesis was mostly centered on genes and ignored the development of the organism, as well as morphological areas of biology.
The extended synthesis is still a work-in-progress, essentially having been launched in 2007 by a paper by Massimo Pigliucci in the journal Evolution, followed by a conference on the subject in 2008 at the Konrad Lorenz Institute for Evolution and Cognition Research. The participants of the conference published a book in 2010 titled Evolution: The Extended Synthesis, which has served as a launching point for work on the extended synthesis. Several of the topics included in the extended synthesis include:
- Showing the importance of the role of prior configurations, genomic structures, and other traits in the organism in generating evolutionary variations.
- Looking at how increasing dimensionality of fitness landscapes affects our view of speciation.
- Examining the role of multilevel selection in the major evolutionary transitions.
- Looking at new types of inheritance, including cultural and epigenetic inheritance.
- Examining the way that organismal development and developmental plasticity channels evolutionary pathways.
- Looking at how organisms modify the environments they belong to through niche construction.
The goal of the 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.
Biologist Eugene Koonin writing on the extended synthesis in 2009 wrote 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."
The Extended Evolutionary Synthesis proposed by Pigliucci and colleagues makes the following predictions:
- change in phenotype can precede change in genotype
- changes in phenotype are predominantly positive, rather than neutral (see: neutral theory of molecular evolution)
- changes in phentotype are induced in many organisms, rather than one organism
- revolutionary change in phenotype can occur through mutation or facilitated variation
- "repeated evolution in isolated populations may be due to convergent selection and/or developmental bias"
- adaptation occur due to natural selection, environmental induction, non-genetic inheritance, learning and cultural transmission (see: Baldwin effect, meme, transgenerational epigenetic inheritance, ecological inheritance, non-Mendelian inheritance)
- rapid evolution can result from the simultaneous induction and natural selection
- biodiversity can be affected by features of developmental systems (evolvability, constraints)
- heritable variation is directed towards variants that are adaptive and integrated with phenotype
- "niche construction will be systematically biased towards environmental changes that are well suited to the constructor's phenotype, or that of its descendants, and enhance the constructor's, or its descendant's, fitness"
- multilevel selection
- kin selection
Biologists disagree on the need for an extended synthesis. Opponents contend that the modern synthesis is able to fully account for the newer observations, while 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.
The ideas of the extended synthesis were positively reviewed by Anya Plutynski for the National Center for Science Education.
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Defend the extended synthesis
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Criticism of the extended synthesis
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- Mayr, Ernst. (2004). 80 years of watching the evolutionary scenery. Science 305: 46-47.
- Merlin, Francesca. (2010). "Evolutionary Chance Mutation: A Defense of the Modern Synthesis’ Consensus View". Philosophy & Theory in Biology 2: 22.
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