(A) Crystal structure of an initiation complex with RRR
discriminator and nontemplate-strand length corresponding to TSS at position 7
(RPo-GGG-7). Left, simulated annealing |Fo-Fc| omit map contoured at
2.3σ and atomic model for interactions of RNAP with DNA
nontemplate-strand. Right, schematic path of DNA. Gray, RNAP; yellow, σ;
purple, discriminator nucleotides; pink, nontemplate-strand nucleotides
downstream of discriminator; green, simulated annealing |Fo-Fc| omit map
contoured at 2.3σ. Red box, third nucleotide of discriminator.
(B) Crystal structure of an initiation complex with YYY
discriminator and nontemplate-strand length corresponding to TSS at position 8
(RPo-CCC-8; symbols as in A).
(C–D) Interpretation: The different position of the third
nucleotide of the discriminator in the structure with a YYY discriminator
increases the distance between the third nucleotide of the discriminator and
downstream duplex DNA, accommodating an additional nucleotide in the
connector.