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. 2026 Feb 13;49(Suppl 2):e20250179. doi: 10.1590/1678-4685-GMB-2025-0179

Table 5- . Core concepts of the Extended Evolutionary Synthesis (EES).

Concept / Mechanism Description Biological Implications Key Authors/Sources
Phenotypic Plasticity Capacity of a genotype to produce different phenotypes in response to environmental cues Allows rapid, reversible adaptation; may precede genetic changes West-Eberhard (2003)
Epigenetic Inheritance Transmission of gene expression patterns via DNA methylation, histone modification, etc. Enables non-genetic adaptation; revives soft inheritance concepts Jablonka and Lamb (2005); Bonduriansky and Day (2009)
Developmental Bias Certain phenotypes are more likely due to constraints or propensities in developmental systems Not all variation is random; evolution is shaped by developmental dynamics Müller (2007); Pigliucci (2007)
Niche Construction Organisms modify their environments in ways that influence their own evolution Feedback loop between organisms and environments; breaks nature/nurture dualism Odling-Smee et al. (2003)
Inclusive Inheritance Broadens heredity to include genetic, epigenetic, ecological, and cultural transmissions Inheritance is multilayered and context-dependent Laland et al. (2015)
Symbiosis and Holobionts Organisms evolve in tight association with symbiotic partners (microbiome, virome, etc.) Challenges individualistic views of evolution; emphasizes community-level selection Gilbert et al. (2012b); Bordenstein and Theis (2015)
Facilitated Variation Conserved core processes allow evolutionary innovation through recombination of modular systems Explains evolvability and the emergence of complexity Gerhart and Kirschner (2007)
Cultural and Behavioral Inheritance Behaviors and learned information passed across generations influencing fitness Particularly important in humans and other social species Laland and Brown (2002)
Reciprocal Causation Evolutionary processes involve feedbacks where cause and effect are not strictly linear Evolution is dynamic, co-constructed, and context-sensitive Laland et al. (2015); Uller and Laland (2019)
Extended Organism Organisms build structures (nests, burrows, webs) that influence selection and are part of phenotype Redefines phenotype to include ecological structures Jablonka and Lamb (2005); Dawkins (2004)
Dissipative Structures Organisms as open, far-from-equilibrium systems that sustain internal order by continuous energy throughput and entropy export Embeds thermodynamic constraints in evolutionary innovation; explains emergence of hierarchical complexity via non-equilibrium dynamics Prigogine and Stengers (1984); Kleidon and Lorenz (2005); Kondepudi et al. (2020)