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5 - Higher-level structures and the recapitulation of metabolic order

Published online by Cambridge University Press:  05 June 2016

Eric Smith
Affiliation:
Tokyo Institute of Technology
Harold J. Morowitz
Affiliation:
George Mason University, Virginia
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Summary

Metabolism exists on Earth today only in a context formed by multiple higher-level structures, which are themselves constructed by living processes. In this chapter we consider four kinds of higher-level living order: ribosomal translation, some broad classes of oligomer catalysts, the tripartite bioenergetic system of redox couples, protons, and phosphate esters, and cellular compartmentalization including the association of cells with genomes. Like metabolism considered in Chapter 4, these higher levels show modular architecture and suggest a history of independent subsystems that were brought together to form extant cells. Many of the module boundaries follow divisions already seen in metabolism. Biosynthetic pathway patterns and a layered structure in the genetic code may reflect layers in the accretion of components of the translation system. The unification of bioenergetics by cells mirrors hierarchy in biochemistry, with a core of redox and thioester activation, and large-scale incorporation of phosphates only later, perhaps enabling the rise of an oligomer world. The cell is not merely one kind of compartment but at least three. The three core functions of cellularization – unification of bioenergetics, catalytic rate enhancement, and homeostatic regulation of the cytosol – may have come at different stages of separation from mineral-hosted environments. The resulting picture of life is of a confederacy of subsystems, which retain some distinct identity even in their current union. During biogenesis these gained autonomy from the environments that drove them into existence by becoming more dependent on each other.

Coupled subsystems and shared patterns

Universal metabolism at the ecosystem level is the chemical source of life, and (as we will argue in Chapter 8) in many respects its informatic foundation as well. However, the distinctive chemistry of living systems only occurs on Earth today in a context of elaborate higher-level structure. Biosynthesis depends in essential ways on catalysis by oligomers synthesized through ribosomal translation, on integrated and regulated bioenergetic systems, and on containment in cells. The maintenance and optimization of all these systems takes place through selection on genes and genomes, which must then be transmitted together during descent to maintain their functionality. These higher-level systems present whole webs of interdependency and entwined chicken-egg paradoxes. To understand the origin of life we cannot only pursue the relations of metabolism to geochemistry.We must also decipher the relations among these higher-level structures into patterns of serial dependency.

Type
Chapter
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The Origin and Nature of Life on Earth
The Emergence of the Fourth Geosphere
, pp. 273 - 339
Publisher: Cambridge University Press
Print publication year: 2016

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