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The Ribosomal “Tree of Life” Cenancestor (aka LUCA or MRCA) can be placed using the echo remaining from the early expansion of the genetic code. • reflects only a single cellular component • no reticulation • only extant lineages • branch length is not proportional to time Deep Paralogs • Gene families evolve by duplication, gene transfer and divergence • For ancient proteins, some of these divergences occurred before the divergence of the three domains of life (MRCA); • Determining function of paralog ancestors could elucidate primordial physiology… however, lack of time machine prevents direct experimental observation. • In some special cases, can be inferred using amino acid composition… Catalytic subunits Non catalytic subunits speciation time gene duplication Gogarten et al, 1989, PNAS 86, 17, 6661–6665 http://www.pnas.org/content/113/32/E4654.full.pdf?with-ds=yes ATPase / ATPsynthase ATP binding Subunits B A A B B V-proteolipid A C N c A-proteolipids A-type ATPase N a b b a N V-type ATPase a C C Halobacterium Methanococcus Methanopyrus b c F-type ATPase F-proteolipid N C 12 proteolipid Ds / 3 catalytic SU = 4H+(Na+) / ATP 6 proteolipid Ds / 3 catalytic SU = 2H+(Na+) / ATP Reversible Enzyme Dedicated Ion Pump Reversible Enzyme Bacteria Eukarya Archaea 12 proteolipid Ds / 3 catalytic SU = 4H+(Na+) / ATP endosymbionts thermophilic mesophilic E D 5 4 ? C B 12 proteolipid Ds / 3 catalytic SU = 4H+(Na+) / ATP Reversible Enzyme A 3 12 proteolipid Ds / 6 catalytic SU = 2 H+(Na+) / ATP 2 Dedicated Ion Pump 1 Evolution of the Ribosome ● ● ● ● “Core” of ribosome consists of RNA + subset of ribosomal proteins universally conserved in all life (~29 proteins) (Harris et al., 2003) mRNA, tRNA, rRNA all crucial components for translation Catalytic peptidyltransferase activity is RNAmediated Likely coevolved with genetic code within an RNA world (Wolf & Koonin, 2007) Rooting the Ribosomal Tree of Life using an Echo from the Early Expansion of the Genetic Code (Fournier and Gogarten, MBE 2010)