The Boggarts of biology: how non-genetic changes influence the genotype
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MINI-REVIEW
The Boggarts of biology: how non‑genetic changes influence the genotype Laasya Samhita1 Received: 30 July 2020 / Revised: 14 September 2020 / Accepted: 15 September 2020 © Springer-Verlag GmbH Germany, part of Springer Nature 2020
Abstract The notion that there is a one–one mapping from genotype to phenotype was overturned a long time ago. Along with genotype and environment, ‘non-genetic changes’ orchestrated by altered RNA and protein molecules also guide the development of phenotype. The idea that there is a route through which changes in phenotype can lead to changes in genotype impinges on several phenomena of molecular, developmental, evolutionary and applied interest. Phenotypic changes that do not alter the underlying DNA sequence have been studied across model systems (eg: DNA and histone modifications, RNA editing, prion formation) and are known to play an important role in short-term adaptation. However, because of their transient nature and unstable inheritance, the role of such changes in long-term evolution has remained controversial. I classify and review three ways in which non-genetic changes can influence genotype and impact cellular fitness across generations, with an emphasis on the enticing idea that they may act as stepping stones for genetic adaptation. I focus on work from microbial systems and attempt to highlight recent experiments and models that bear on this idea. Overall, I review evidence which suggests that non-genetic changes can impact phenotype via their influence on the genotype, and thus play a role in evolutionary change. Keywords Non-genetic change · Translation errors · Phenotypic variability · Adaptation · Evolution
Introduction Heritable phenotypic variability, i.e., phenotypic changes that are faithfully passed on from generation to generation, lies at the heart of Darwinian evolution. Typically, heritable variation stems from differences in the genetic material (usually DNA) of the organism. In contrast to this, recent findings show that non-DNA-based changes can also lead to phenotypic variability, impact fitness, and be inherited (reviewed in Bonduriansky and Day 2009; Jablonka and Raz 2009; Ribas de Pouplana et al. 2014, Charlesworth et al. 2017). Many terms have been coined to refer to non-DNA based change; in fact the whole field of ‘epigenetics’ is devoted to their study. Perhaps the most familiar examples under this category are those of DNA and histone modifications. These topics, along with RNA editing, have been Communicated by M. Kupiec. * Laasya Samhita [email protected]; [email protected] 1
National Centre for Biological Sciences, Tata Institute of Fundamental Research, Bangalore, India
extensively reviewed elsewhere (Gott and Emeson 2000; Jablonka and Raz 2009; Charlesworth et al. 2017). For the purpose of this article and to avoid confusion, I use the term ‘non-genetic changes’ to mean changes that impact phenotype without altering the primary DNA sequence. At a molecular level, RNA and protein alterations underlie nearly all such cha
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