TABLE 1.

+1 frameshifts induced by altered tRNAs

tRNA or stress	Alteration in tRNA	Site of frameshifta	Expt type	Reference(s)
S. enterica tRNAGGGPro	Base substitutions outside and in anticodon; tRNA aminoacylated as in WT	P site, by WT tRNAcmo5UGGPro	In vivo	258, 259, 295
S. enterica tRNACGGPro	Insertion of an extra nucleotide in anticodon	P site, by either mutated tRNACGGPro or WT tRNAcmo5UGGPro	In vivo	259
Yeast tRNAIGGPro	Insertion of an extra nucleotide in anticodon	P site	In vivo	259
Yeast tRNACCCGly	Insertion of an extra nucleotide in anticodon	P site	In vivo	259
S. enterica tRNAmnm5UCUArg	Alterations in structure of tRNAmnm5UCUArg reduce concn of aminoacylated tRNAmnm5UCUArg, which in turn causes frameshift at next upstream codon	P site	In vivo	180
Starvation	Amino acid limitation	P site	In vivo	107
Third-position-mismatched tRNAs	Lack of cognate tRNAs induces +1 frameshift by near-matched tRNA; i.e., imbalance of tRNAs causes +1 frameshift	P site	In vivo	304
Imbalance of tRNAcmo5UGGPro or tRNAmnm5UCUArg	Overexpression of tRNAcmo5UGGPro induces +1 frameshift; +1 frameshift at tandem AGG-AGG caused by limitation of tRNAmnm5UCUArg	P site	In vivo	229, 291
Third-position-mismatched tRNAs	Several third-position-mismatched tRNAs in P site induce frameshift	P site	In vivo	304
tRNA modifications	Various deficiencies in tRNA modification cause +1 or −1 frameshifts in E. coli, S. enterica, or yeast	In some cases shown to occur in P site; in other cases not addressed	In vivo	40, 120, 176, 188, 320, 325
E. coli tRNAmnm5UCCGly	Base substitutions in position 34 and outside anticodon	P site	In vivo	131
E. coli tRNAcmo5UACVal	Insertion of either A or U in anticodon between cmo5U34 and A35	P site	In vivo	235
S. enterica tRNAGGUThr	Insertion of an extra nucleotide in 5′ part of the anticodon and “pushing” U33 into wobble position, resulting in 5′-UUGG-3′ anticodon	Possible P site (see text)	In vivo	45, 46
S. enterica tRNAcmnm5s2UUGGln	Overexpression of tRNAcmnm5s2UUGGln with an inserted U between cmnm5s2U34 and U35	Not experimentally addressed	In vivo	228
S. enterica tRNAmnm5s2UUULys	Overexpression of tRNAmnm5s2UUULys with an inserted U between mnm5s2U34 and U35	Not experimentally addressed	In vivo	228
E. coli tRNAICGArg	Changes in or outside anticodon	Not experimentally addressed	In vivo	317
E. coli tRNACUALeu	Insertions of an extra nucleotide in anticodon and A26G base substitution	Not experimentally addressed	In vivo	211
E. coli tRNAcmo5UACVal	Insertion of an extra nucleotide (C32.1) and G42A present on plasmid	Not experimentally addressed	In vivo	230
E. coli tRNAcmo5UACVal	Base substitutions (C74A or C74G) in the CCA terminus	Not experimentally addressed	In vivo	239
Mitochondrial tRNAUGASer	U42C and lack of Ψ27.	Not experimentally addressed	In vivo	143
tRNAAla	Insertion of an extra nucleotide in anticodon; in vitro experiments	Not experimentally addressed	In vitro	326
tRNAPhe	Insertion of CCCG as anticodon in body of unmodified tRNAPhe; in vitro experiments	Not experimentally addressed	In vitro	253

^aThe site of frameshifting was determined as described in the text under Tests of P-Site and E-Site Roles: Genetic Evidence for E-Site Codon-Anticodon Pairing.