By Albert Neuberger, L. L. M. Van Deenen, Giorgio Semenza
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Extra info for A History of Biochemistry: Selected Topics in the History of Biochemistry Personal Recollections. II.
Greenberg's report  on the development of a cell-free system was followed in time with other major contributions. One such contribution was the finding that inosinic acid is the first compound formed with a completed purine ring and that it is the precursor of inosine and hypoxanthine, and not vice versa . These experiments demonstrated that the elements of ribose phosphate became attached to a purine precursor prior to final completion of the ring (Fig. 2). This finding was followed within a short time by A BACKWARD GLANCE 23 GLUTAMINE GLUTAMATE +H2O +HP2O"; 03POCH io^f H OH O;POCH.
For this reason DNA-RNA competitionhybridization experiments were undertaken by Grasso  in this laboratory and by Salser et al.  to study the synthesis of early T4 mRNAs under a variety of conditions, for example, during inhibition of protein synthesis. When chloramphenicol, an agent that effectively inhibits protein synthesis in prokaryotes, is included at the time of infection of E. coli B with T4 bacteriophage, a class of mRNAs is formed that may represent up to 30% of the total early species.
With the discovery of the set of enzymes concerned with the synthesis of phage DNA mentioned above, we began a collaborative effort aimed at the identification of the functions or activities missing in the conditional lethal amber mutants. The first to be established was the assigment of dCMP hydroxymethylase to gene 42 . DNA polymerase was also lacking in these extracts, but the specific mutant of gene 42 studied, am N122, was later shown to contain a double mutation. The genetic contribution of both the bacteriophage and the host to the production of mutant enzyme was next explored by Wiberg and by Dirksen and Hutson, respectively.