Toward understanding of evolutionary constraints: experimental and theoretical approaches

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Toward understanding of evolutionary constraints: experimental and theoretical approaches Chikara Furusawa 1,2

&

Naoki Irie 2,3

Received: 12 May 2020 / Accepted: 11 June 2020 # International Union for Pure and Applied Biophysics (IUPAB) and Springer-Verlag GmbH Germany, part of Springer Nature 2020

Abstract Although organisms have diversified remarkably through evolution, they do not exhibit unlimited variability. During evolution, the phenotypic changes do not occur at random; instead, they are directional and restricted by the constraints imposed on them. Despite the perceived importance of characterizing the unevenness of these changes, studies on evolutionary constraints have been primarily qualitative in nature. In this review, we focus on the recent studies of evolutionary constraints, which are based on the quantification of high-dimensional phenotypic and genotypic data. Furthermore, we present a theoretical analysis that enables us to predict evolutionary constraints on the basis of phenotypic fluctuation, modeled on the fluctuation–response relationship in statistical physics. The review lays emphasis on the tight interactions between experimental and theoretical analyses in evolutionary biology that will contribute to a better understanding of evolutionary constraints. Keywords Evolutionary constraints . Microbial laboratory evolution . Developmental hourglass model . Evolutionary fluctuation–response relationship

Introduction Although organisms have undergone remarkable diversification during evolution, it has not occurred in entirely random directions. Previous studies have shown unevenness and directionality in the evolutionary changes and phenotypic variations (Arnold 1992; Smith et al. 1985). For example, all vertebrates have a maximum of two pairs of limbs, and all insects have the same basic body plan (Urry et al. 2016). These limits of diversity have often been attributed to the limited potential for diversification or constraints on the production of variable phenotypes; however, the exact mechanism remains largely unclear.

* Chikara Furusawa [email protected] * Naoki Irie [email protected] 1

Center for Biosystems Dynamics Research, RIKEN, 6-2-3 Furuedai, Suita, Osaka 565-0874, Japan

2

Universal Biology Institute, The University of Tokyo, 7-3-1 Hongo, Bunkyo-ku, Tokyo 113-0033, Japan

3

Department of Biological Sciences, The University of Tokyo, 7-3-1 Hongo, Bunkyo-ku, Tokyo 113-0033, Japan

The theoretical approach, which is based on the concept of statistical physics, contributed significantly toward formulating the constraints, leading to tight collaborations between theoretical and experimental studies (Kaneko and Furusawa 2018; Sato et al. 2003). Technological innovations also enabled the quantitative analyses of high-dimensional phenotypic and genotypic data, allowing scientists to experimentally test the theoretical frameworks. The present review highlights the contribution of quantitative and theoretical analyses of evolutionary dynamics toward the underst