Transcription of DNA is a vital process for gene expression and the regulation of proteins in prokaryotic and eukaryotic cells.

The fundamentals of transcription are the processes of binding, initiation, elongation and termination, which occur in both eukaryotes and prokaryotes. However, as eukaryotic DNA is more complexly arranged in chromosomes, prokaryotic DNA is circular and not contained within a nucleus, being just one of the many reasons for differences in the transcription process. Firstly, in prokaryotes and eukaryotes the gene to be transcribed (coding strand) has a promoter region, which is the start point for the enzyme RNA polymerase to first bind to. Generally in prokaryotes, where transcription takes place in the cell cytoplasm, the promoter region falls between DNA sequence positions -35 and -10, known as the Pribnow box, it has been suggested by Beebee & Burke (1988) that promoter activity in prokaryotes is dependent on a maximum distance of 17 nucleotide bases between these sequences. A holoenzyme containing the RNA polymerase, sigma factor and other and subunits, uses the sigma factor to aid the binding of RNA polymerase to the promoter region forming a closed promoter complex. A difference in the control of transcription is that prokaryotes have a cluster of genes that are linked together in an operon, where an activating signal will result in the transcription of all the genes of the operon as one single polycistronic mRNA molecule (Latchman, 1995). This is very different from the control of transcription in eukaryotes as only individual genes are transcribed into individual mRNA molecules. Even though eukaryotes and prokaryotes both use RNA polymerase to bind to the promoter region on the gene, the difference lies where eukaryotes have 3 types of RNA polymerase (I, II, III) to accommodate the different sizes of rRNA, protein coding RNA and tRNA (Beebee & Burke, 1988). Eukaryotes also require specific transcription factors that bind to the promoter region to form a transcription initiation complex. The TATA box is a promoter region that uses the transcription factor TBP (TATA-binding protein) to bind to it, which allows other factors such as transcription factor D to bind to the promoter region as well, the transcription initiation complex is formed when RNA polymerase and all the factors are bound to the promoter region. Therefore, protein-protein interactions also have an important part in the initiation of transcription in eukaryotes, because some transcription factors may bind directly to DNA while others may bind to other transcription factors to initiate transcription (Hardin et al, 2010). In consequence, once initiation takes place, the double helix of DNA is unwound as RNA

polymerase will copy the DNA template strand into a complementary RNA strand in the 5 to 3 direction, common for both prokaryotes and eukaryotes. In the newly formed RNA strand, Thymine (T) is replaced by Uracil (U), as the complementary binding of nucleotide base pairs takes place (Adenine with Uracil, and Cytosine with Guanine), the continual forming of the RNA is known as elongation. For eukaryotes, elongation requires the breakdown of the nucleosomes that are ahead of the moving RNA molecule, complex proteins will perform this and also repair the nucleosomes once the DNA is transcribed. During elongation, topoisomerase enzymes work on the DNA to control the supercoiling of it in both prokaryotes and eukaryotes. RNA proofreading can correct mistakes such as noncomplementary base pairs being added to the sequence, in which the nucleotide will remove itself (Hardin et al, 2010). Elongation factors like elongin will help prevent pausing of the elongation process, making it more sufficient (White, 2001). The termination of transcription is probably the most distinct between the two kingdoms, it will happen once the RNA reaches the termination signal. In prokaryotes there are two types of termination signals, reliant upon whether or not the RNA molecule forms a hairpin loop. If the RNA is rich in GC base pairs and U nucleotide bases at the 3 end, then this loop is formed simultaneously causing the RNA to pull away from the DNA, and the bonds between the U on the RNA and A on the DNA template strand are broken (Hardin et al, 2010). However if this hairpin loop is not formed a (Rho) factor is required. The factor will use ATP to unwind the RNA from the DNA template once it has been bound to a specific termination sequence. In eukaryotes, the termination process is achieved by a cleavage formation in the transcript. Cleavage factors such as CPSF (Cleavage and Polyadenylation Specificity Factor) binds the AAUAAA sequence, the cleavage is formed between this polyadenylation signal and a number of U nucleotides that are downstream (White, 2001). Hence, approximately 10-35 nucleotides after RNA Polymerase II transcribes this signal, proteins will cut off the mRNA to release it for processing, where the 5 end receives a different form of Guanine bases (Cain et al, 2011). Finally, the 3 end receives adenylate to form a poly(A) tail of approximately 250 nucleotides (White, 2001). To conclude, transcription is more complex in eukaryotes with the addition of forming an RNA cleavage to terminate the process and then further RNA processing. This may be because prokaryotes being unicellular have a less complex existence, so the produced RNA molecule has less phenotypes to produce than does eukaryotic RNA. References

Latchman, D. (1995) Gene Regulation: A eukaryotic perspective. 2nd edn. London: Chapman & Hall. Beebee, T. & Burke, J. (1988) Gene Structure And Transcription. Oxford: IRL Press Limited. White, R.J. (2001) GeneTranscription. Oxford: Blackwell Science Ltd. Cain, M.L. Jackson, R. B. Minorsky, P.V. Reece, B.J. Urry, L.A. Wasserman, S.A. (2011) Campbell Biology. Global Edition. San Francisco: Pearson Education. White, R.J. (2001) GeneTranscription. Oxford: Blackwell Science Ltd. Bertoni, G. Hardin, J. Kleinsmith, L.J. (2012) Beckers World of the Cell. 8th edn. San Francisco: Pearson Education, Pearson Benjamin Cummings.