The amber mutants discovered by Charles Steinberg in collaboration with Richard Epstein provided a unique opportunity to study the function of virtually all the genes of phage T4 that are required for growth of the phage under laboratory conditions. The circumstances under which the amber mutants were discovered was described by Edgar in a retrospective article.[9] Soon after their discovery, the phage amber mutants, initially characterized by Steinberg and Epstein, were made available to many other investigators. These mutants were used to obtain important insights into fundamental aspects of biology. The amber mutants contributed to our understanding of the functions and interactions of the proteins employed in DNA replication, DNA repair, genetic recombination and the assembly of viruses from their molecular components.