MOLECULAR BASIS OF INHERITANCE
(28S, 18S, and 5.8S), whereas the RNA polymerase III is responsible
for transcription of tRNA, 5srRNA, and snRNAs (small nuclear
RNAs). The RNA polymerase II transcribes precursor of mRNA, the
heterogeneous nuclear RNA (hnRNA).
(ii) The second complexity is that the primary transcripts contain both
the exons and the introns and are non-functional. Hence, it is
subjected to a process called splicing where the introns are removed
and exons are joined in a defined order. hnRNA undergoes
additional processing called as capping and tailing. In capping an
unusual nucleotide (methyl guanosine triphosphate) is added to
the 5' -end of hnRNA. In tailing, adenylate residues (200-300) are
added at 3' -end in a template independent manner. It is the fully
processed hnRNA, now called mRNA, that is transported out of the
nucleus for translation (Figure 6.11).
The significance of such complexities is now beginning to be
understood. The split-gene arrangements represent probably an ancient
feature of the genome. The presence of introns is reminiscent of antiquity,
and the process of splicing represents the dominance of RNA-world. In
recent times, the understanding of RNA and RNA-dependent processes
in the living system have assumed more importance.
6.6 GENETIC CODE
During replication and transcription a nucleic acid was copied to form
another nucleic acid. Hence, these processes are easy to conceptualise
on the basis of complementarity. The process of translation requires
transfer of genetic information from a polymer of nucleotides to synthesise
a polymer of amino acids. Neither does any complementarity exist between
nucleotides and amino acids, nor could any be drawn theoretically. There
existed ample evidences, though, to support the notion that change in
nucleic acids (genetic material) were responsible for change in amino acids
in proteins. This led to the proposition of a genetic code that could direct
the sequence of amino acids during synthesis of proteins.
If determining the biochemical nature of genetic material and the
structure of DNA was very exciting, the proposition and deciphering of
genetic code were most challenging. In a very true sense, it required
involvement of scientists from several disciplines – physicists, organic
chemists, biochemists and geneticists. It was George Gamow, a physicist,
who argued that since there are only 4 bases and if they have to code for
20 amino acids, the code should constitute a combination of bases. He 111
suggested that in order to code for all the 20 amino acids, the code should
be made up of three nucleotides. This was a very bold proposition, because
a permutation combination of 43 (4 × 4 × 4) would generate 64 codons;
generating many more codons than required.
Providing proof that the codon was a triplet, was a more daunting
task. The chemical method developed by Har Gobind Khorana was
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instrumental in synthesising RNA molecules with defined combinations
of bases (homopolymers and copolymers). Marshall Nirenberg’s cell-free
system for protein synthesis finally helped the code to be deciphered.
Severo Ochoa enzyme (polynucleotide phosphorylase) was also helpful
in polymerising RNA with defined sequences in a template independent
manner (enzymatic synthesis of RNA). Finally a checker-board for genetic
code was prepared which is given in Table 6.1.
Table 6.1: The Codons for the Various Amino Acids
The salient features of genetic code are as follows:
(i) The codon is triplet. 61 codons code for amino acids and 3 codons do
not code for any amino acids, hence they function as stop codons.
(ii) Some amino acids are coded by more than one codon, hence
the code is degenerate.
(iii) The codon is read in mRNA in a contiguous fashion. There are
no punctuations.
(iv) The code is nearly universal: for example, from bacteria to human
UUU would code for Phenylalanine (phe). Some exceptions to this
rule have been found in mitochondrial codons, and in some
protozoans.
(v) AUG has dual functions. It codes for Methionine (met) , and it
112 also act as initiator codon.
(vi) UAA, UAG, UGA are stop terminator codons.
If following is the sequence of nucleotides in mRNA, predict the
sequence of amino acid coded by it (take help of the checkerboard):
-AUG UUU UUC UUC UUU UUU UUC-
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