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Transcription -

A gene is transcribed into RNA by a DNA-dependent RNA polymerase, which copies a single stranded DNA template using the four ribonucleotide triphosphates as substrates. Plant cells, in common with animal cells, contain three types of RNA polymerase which can be distinguished on the basis of physical properties, localization and function. RNA polymerase II is responsible for the synthesis of mRNA precursors and is located in the nucleoplasm. RNA polymerases have been isolated and characterized from a wide range of plant species; they carry out the first vital step in gene expression.

RNA polymerase I synthesizes only ribosomal RNA; RNA polymerase III synthesizes tRNAs and some other small RNAs. RNA polymerase, II is responsible for the synthesis of RNA from structural genes. Promoter regions for RNA polymerase II have been estimated from a comparison of the base sequences upstream from many different eukaryotic genes. There are two "consensus" regions that are good candidates for promoters.

RNA polymerase binds to a promoter region in the 5' flanking sequence of the gene, initiates a mRNA chain and then copies through the gene until it reaches a termination signal. Eukaryotic promoters are distinct from those in prokaryotes and are not efficiently recognized by prokaryote polymerase.

Control of transcriptions is a vital element in the overall control of gene expression but is very poorly understood in plants. Studies have been carried out to look at overall levels and activities of the various types of polymerases. Guilfoyle and Malcolm (1980) found that during soybean germination the activities of polymerase I and III increased rapidly after 6 hours, whereas the increase in polymerase II was much less dramatic. An increase in poly­merase activity in etiolated seedlings treated with a growth regulator, auxin was also reported. During germination polymerase II underwent a change in size of its largest subunit to convert the A-form of the enzyme to the B­ form.

Concurrent with this change, the polymerase II activity in the seedlings increased dramatically although the amount of protein remained constant, leading to the hypothesis that the A-form must be converted to the active B-form at the start of germination for transcription to take place.

Template availability has been studied in a few cases but is very dif­ficult to measure and interpret. Some increase in overall template activity can be measured after treating tissues with auxins or gibberellins. Tem­plate availability has been linked in some cases to the types of non histone chromosomal proteins associated with the DNA.

In animal systems control of transcriptions has been shown to involve changes in chromatin structure, methylation of cytosine residues of the DNA and interaction of specific regulatory molecules with the chromatin. Although it is thought likely that such phenomena may also be involved in plant cells, as yet there is little direct supportive evidence for this supposition.