The Emergence of Modularity in Biological Systems
Dirk M. Lorenz, Alice Jeng, and Michael W. Deem

TL;DR
This review explores how modularity and hierarchy naturally emerge in biological systems through symmetry-breaking phase transitions, supported by diverse examples from molecular to ecological levels.
Contribution
It synthesizes theories and experimental evidence explaining the spontaneous organization of biological modularity as a phase transition phenomenon.
Findings
Modularity arises as a symmetry-breaking phase transition.
Experimental evidence supports modularity as an emergent property.
Examples span from protein structures to ecological networks.
Abstract
In this review, we discuss modularity and hierarchy in biological systems. We review examples from protein structure, genetics, and biological networks of modular partitioning of the geometry of biological space. We review theories to explain modular organization of biology, with a focus on explaining how biology may spontaneously organize to a structured form. That is, we seek to explain how biology nucleated from among the many possibilities in chemistry. The emergence of modular organization of biological structure will be described as a symmetry-breaking phase transition, with modularity as the order parameter. Experimental support for this description will be reviewed. Examples will be presented from pathogen structure, metabolic networks, gene networks, and protein-protein interaction networks. Additional examples will be presented from ecological food networks, developmental…
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