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. 2022 Mar 4;90(2):166–175. doi: 10.1007/s00239-022-10049-1

Table 1.

Exaptation of proteins and RNAs

Ancestral functiona Exapted function(s)
Protein
Cytosolic ribosomal proteins (Wang et al. 2015; Lu et al. 2015) Structure and assembly of the ribosome In transcription, cell growth and proliferation, apoptosis, mRNA splicing, DNA repair, cellular development, and cellular differentiation
Ribonuclease III (Petrov et al. 2019) Cleavage of double-stranded RNA As a mitochondrial ribosomal protein
MutT (Petrov et al. 2019) Repair of DNA containing 8-oxoguanine As a mitochondrial ribosomal protein
Aminoacyl-tRNA synthetases (Guo and Schimmel 2013) Covalently link amino acids to their cognate tRNAs In metabolism, development, angiogenesis, tumorigenesis, immune response, neuronal function, and inflammation
Glyceraldehyde-3-phosphate dehydrogenase (GAPDH) (Singh and Bhalla 2020) Catalyzes the sixth step of glycolysis, the conversion of d-glyceraldehyde 3-phosphate to 3-phospho-d-glyceroyl phosphate In apoptosis, iron transport, membrane fusion, transcriptional regulation, vesicle transport, and cellular response to oxidative stress and hypoxia
Argininosuccinate lyase (Piatigorsky 2003; Gavelis et al. 2017) Catalyzes the fourth step of the urea cycle and is involved in the biosynthesis of arginine Light focusing δ-crystallins in birds and reptiles
Lactate dehydrogenase (Hendriks et al. 1988) Catalyzes the reversible conversion of pyruvate to lactate and of NADH to NAD+ Light focusing ε-crystallins in birds and reptiles
Pancreatic trypsinogen (Chen et al. 1997) A zymogen of trypsin, a digestive protease An antifreeze protein in cold water fishes
Transposase (Fugmann 2010) Cut-and-paste transposition A V(D)J recombinase, which rearranges immunity-related genes
Protein of an endogenous retrovirus (Cornelis et al. 2015) Retrovirus envelope formation A mediator of placentation in mammals
RNA
tRNAs (Luchetti and Mantovani 2013; Robeck et al. 2016) Translational adaptors BC1 and many additional short interspersed nuclear elements (SINEs) in animals with many functions
tRNAval and tRNAPhe (Greber et al. 2014; Rorbach et al. 2016; Brown et al. 2017) Translational adaptors Replacements for 5S rRNA in mammalian mitochondrial ribosomes
tRNA fragments (Schorn and Martienssen 2018; Magee and Rigoutsos 2020) Translational adaptors Guides to silencing of mRNAs by binding to argonaute and PIWI proteins, inhibitors of retrotransposition by binding to the primer binding site of LTR-retrotransposons, inhibitors of replication of retroviruses and LTR-retroelements, inhibitors of translation, and mediators of epigenetics
Dormant transposon (Ellis et al. 2018; Brosius 2019) Transposon Post-transcriptional regulation

aIn several cases the characterization of ancestral versus exapted/adapted functions (i.e., the polarity) is tentative. Ancestry is relative to more recent exapted functions and should not be interpreted to imply ultimate ancestry

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