Protein synthesis in ribosomes requires two kinds of tRNAs:
initiation and elongation. The former initiates the process
(formylmethionine tRNA in prokaryotes and special methionine
tRNA in eukaryotes). The latter participates in the synthesis
proper, recognizing the sense codons. Synthesis is also assisted
by special proteins: initiation, elongation, and termination
factors. The termination factors are necessary to recognize
stop codons (UAG, UGA, and UAA) and to release the complete
protein chain from the elongation tRNA preceding a stop codon.
No termination tRNA capable of recognizing stop codons by their
anticodons is known. The termination factors are thought to
do this. In the large ribosomal RNA, we found two sites that,
like tRNAs, contain the anticodon hairpin but with triplets
complementary to stop codons. One site is hairpin 69 from domain
IV; the other site is hairpin 89, domain V. By analogy, we call
them termination tRNAs: Ter-tRNA1 and Ter-tRNA2, respectively,
even though they transport no amino acids, and suggest that
they directly pair to stop codons. The termination factors only
aid in this recognition, making it specific and reliable. A
strong argument in favor of our hypothesis comes from vertebrate
mitochondria. They are known to acquire two new stop codons,
AGA and AGG. In the standard code, these are two out of six
arginine codons. We revealed that the corresponding anticodons,
UCU and CCU, have evolved in Ter-tRNA1 of these mitochondria.