The effect of DNA supercoiling on gene expression is dependent not onl
y on specific genes but also on the sequence context of the genes. Thi
s position-dependent supercoiling effect on gene activation is best il
lustrated in the study of the suppression of the leu-500 mutation of t
he leuABCD operon in a Salmonella typhimurium topA mutant. In this com
munication, we report a novel promoter relay mechanism whereby several
genes are sequentially expressed in a position-dependent manner: the
ilvIH promoter (pilvIH) activates a cryptic leu0 promoter (pleu0) loca
ted between the two divergently arrayed ilvIH and leu-500 promoters. B
oth the cis-acting pleu0 activity and the trans-acting Leu0 protein ar
e necessary for subsequent activation of the leu-500 promoter (pleu-50
0). Furthermore, pleu0 can be functionally replaced with the inducible
tac promoter (ptac) for leu-500 activation, suggesting that transcrip
tion-driven DNA supercoiling underlies the relay mechanism. This is th
e first example of several related genes communicating via a promoter
relay mechanism for their coordinated expression.