Balancing Robustness and Evolvability.docVIP

Essay Balancing Robustness and Evolvability * Richard E. Lenski , Jeffrey E. Barrick, Charles Ofria ecosystem might be robust if it resists the extinction of some keystone species or, if extinction does occur, because surviving species can compensate over physiological, demographic, or evolutionary time scales. O One important question is ne of the most important features of biology is the ability of organisms to persist whether there exists a single unifying mathematical framework that can encompass such diverse examples of biological robustness. Might new insights come from such a conceptual uni?cation, or will future understanding require detailed analyses of speci?c cases? Across the different scales, recurring mechanisms for achieving robustness—including redundancy of component parts and negative feedbacks—might serve as organizing principles. Yet, similarities in mechanism could mask important differences in the evolutionary origins of those mechanisms. At the level of genes in genomes or of cells in multicellular organisms, it is reasonable to suggest that redundancy evolved by natural selection to maintain some functional capacity in the face of perturbation [4]. But whereas species redundancy could also be critical for robustness of ecosystem functions, differences in redundancy might be an emergent property rather than an ecosystem-level adaptation, because selection generally acts at lower levels (but see [5] for another view). in the face of changing conditions. Consider the remarkable fact that every organism alive today is the product of billions of generations in which its progenitors, without fail, managed to produce progeny that survived to reproduce. To achieve this consistency, organisms must have a balance between robustness and evolvability, that is, between resisting and allowing change in their own internal states [1–3]. Moreover, they must achieve this balance on multiple time scales, including physiological re