, Volume 11, Issue 2, pp 263–285 | Cite as

Macroevolutionary Freezing and the Janusian Nature of Evolvability: Is the Evolution (of Profound Biological Novelty) Going to End?

  • Jan Toman
  • Jaroslav FlegrEmail author
Original Paper


In a macroevolutionary timescale, evolvability itself evolves. Lineages are sorted based on their ability to generate adaptive novelties, which leads to the optimization of their genotype-phenotype map. The system of translation of genetic or epigenetic changes to the phenotype may reach significant horizontal and vertical complexity, and may even exhibit certain aspects of learning behaviour. This continuously evolving semiotic system probably enables the origin of complex yet functional and internally compatible adaptations. However, it also has a second, “darker”, side. As was pointed out by several authors, the same process gradually reduces the probability of the origination of significant evolutionary novelties. In a similar way to the evolution of societies, teachings, or languages, in which the growing number of internal linkages gradually solidifies their overall structure and the structure or interpretation of their constitutive elements, the evolutionary potential of lineages decreases during biological evolution. Possible adaptations become limited to small “peripheral” modifications. According to the Frozen Evolution theory, some of the proximate causes of this “macroevolutionary freezing” are more pronounced or present exclusively in sexual lineages. Sorting based on the highest (remaining) evolvability probably leads to the establishment of certain structural features of complex organisms, e.g. the modular character of their development and morphology. However, modules also “macroevolutionary freeze” whereas the hypothetical “thawing” of modules or their novel adaptive combinations becomes rarer and rarer. Some possible ways out of this dead end include the rearrangement of individual development, e.g. neoteny, radical simplification, i.e. sacculinization, and transition to a higher level of organization, e.g. symbiosis or symbiogenesis. The evolution of evolvability is essentially a biosemiotic process situated at the intersection of the genocentric modern synthesis and the evo-devo-centric extended synthesis. Therefore, evolvability may eventually connect these three not necessarily contradictory approaches.


Evolvability Evolutionary theory Extended synthesis Macroevolution Stability-based sorting Frozen evolution theory 



We thank Charlie Lotterman for the final revisions of our text.


This work was supported by the Grant Agency of the Charles University in Prague (project no: 578416); and the Charles University Research Centre (UNCE 204004). The funding sources had no role in study design, the collection, analysis and interpretation of data, the writing of the report and in the decision to submit the article for publication.

Compliance with Ethical Standards

Conflict of Interest

The authors declare that they have no conflict of interest.


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Authors and Affiliations

  1. 1.Laboratory of Evolutionary Biology, Department of Philosophy and History of Sciences, Faculty of ScienceCharles University in PraguePrague 2Czech Republic

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