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Translation increases the functional diversity of genes by an estimated 6–9 orders of magnitude by converting the linear succession of bases or triplet codons into three-dimensional structures with extraordinary differential binding affinities. The origin of processive codon-dependent peptide bond formation thus provided an extraordinary increase in diversity and precision for nanoscale chemical engineering and control. The proteome is thus, arguably, nature's most majestic invention. Understanding how this invention came about is one of the most significant challenges now facing biology. The past decade has uncovered plausible answers to four questions associated with what enabled biological phenotypes to first separate from genotypes and framed an experimental basis for addressing a fifth. I summarize these developments in this chapter and assess how they help place previous contributions to the evolutionary pre-history of the proteome on a sound experimental footing.

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