Abstract
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.
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Notes
This is not to say evolutionary lineages or the whole biosphere live or learn in the same way individual organisms do. However, in order to avoid any anthropomorphic tone, it would be necessary to discuss these aspects of evolution of evolvability purely in terms of mutually responsive interactions. This would make the exposition less clear and possible analogies with individual learning less obvious. Therefore, we (as numerous authors before us) stuck with the term “learning”, which, however, should not be thought as identical to individual learning but only analogous with it some degree (see, e.g. Davies 2014; Watson et al. 2014, 2016; Lindholm 2015; or Watson and Szathmary 2016 for more detail).
It remains an open question whether all forms of learning and similar processes in living nature are based on the same principle – “domesticated” natural selection. Broad array of processes, e.g. the affinity maturation of antibodies in our immune system (Manser 1990) or “testing” of hypotheses about outside world that define Dennett’s (1995) Popperian organisms, seems to support this possibility.
FPT is, in some regards, close to older punctuational theories of evolution (for systematic review, see Flegr 2013). However, it is based on the existence of alleles with frequency dependent effects on fitness rather than on the existence of epistasis and offers the most complex scenario of related events.
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We thank Charlie Lotterman for the final revisions of our text.
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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.
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Toman, J., Flegr, J. Macroevolutionary Freezing and the Janusian Nature of Evolvability: Is the Evolution (of Profound Biological Novelty) Going to End?. Biosemiotics 11, 263–285 (2018). https://doi.org/10.1007/s12304-018-9326-y
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DOI: https://doi.org/10.1007/s12304-018-9326-y