SUMMARY
The first part of this review contains an overview of the various
contributions and models relating to the control of rRNA synthesis
reported over the last 45 years. The second part describes a systems
biology approach to identify the factors and effectors that control the
interactions between RNA polymerase and rRNA (rrn) promoters
of Escherichia coli bacteria during exponential growth in
different media. This analysis is based on measurements of absolute
rrn promoter activities as transcripts per minute per promoter
in bacterial strains either deficient or proficient in the synthesis of
the factor Fis and/or the effector ppGpp. These absolute promoter
activities are evaluated in terms of rrn promoter strength
(V
max/Km
) and free RNA
polymerase concentrations. Three major conclusions emerge from this
evaluation. First, the rrn promoters are not saturated with
RNA polymerase. As a consequence, changes in the concentration of free
RNA polymerase contribute to changes in rrn promoter
activities. Second, rrn P2 promoter strength is not
specifically regulated during exponential growth at different rates;
its activity changes only when the concentration of free RNA polymerase
changes. Third, the effector ppGpp reduces the strength of the
rrn P1 promoter both directly and indirectly by reducing
synthesis of the stimulating factor Fis. This control of rrn
P1 promoter strength forms part of a larger feedback loop that adjusts
the synthesis of ribosomes to the availability of amino acids via amino
acid-dependent control of ppGpp
accumulation.