Early Transcription of Genes N and Cro
Within a minute after injection
into the host, the complementary ends of the lambda DNA anneal to each other
and are covalently joined by the DNA ligase of the host. Supercoiling of the
circle by DNA gyrase is then possible, and the DNA rapidly acquires the
superhelical density necessary for maximum activity of the phage early
promoters pL and pR. These are located on
either side of CI.
As on all
promoters, initiation of transcription from the promoters pLand pRis
followed by the dissociation of the sigma subunit of RNApolymerase after the
polynucleotide chains have reached a length of 6 to 15 nucleotides. Then a host
protein, the NusA product, binds to RNA polymerase. Nus stands for N
utilization substance. Most likely it binds in the site previously occupied by
the sigma subunit because only one or the other of these proteins can bind to
RNA polymerase at once.
The RNA polymerase molecules elongate past the
point where the sigma and NusA subunits exchange and they transcribe through
the N
gene on the left and the cro gene on the right. This is all that happens at this stage of
the lambda infection, for just beyond these genes the RNA
Figure
14.4 The sigma subunit dissociates
from RNA polymerase and is re-placed by the NusA protein. At the terminator,
the rho protein interacts with the complex and releases RNA and RNA polymerase.
polymerase
pauses, assisted by the NusA protein, and the host transcrip-tion terminator,
rho, releases the RNA chain and RNA polymerase from the DNA (Fig. 14.4). This
pausing and termination occurs at the first transcription termination sites in
the pL and pR operons, tL1 and tR1. Additional terminators with slightly different
properties, tL2 and tR2, lie farther downstream
in both operons. As a result of tL1
and tR1, the first stage
of lambda vegetative growth is confined to the accumulation of N and Cro
proteins.
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