The discovery of bacterial viruses (bacteriophages or phages) in 1941 opened a new era in the study of the genetics of prokaryotic organisms. Although they were disappointing in the original hope that they could be used to fight bacterial infections, phages served during the 1950s as vehicles for genetic analysis of bacteria. Unlike viruses that infect plant or animal cells, phages can relatively easily be analyzed in their host cells. Names associated with phage analysis are Max Delbrück, Salvador Luria, and Alfred D. Hershey (the “phage group,” see: Cairns et al., 1966).

A. Attachment of a bacteriophage

Phages consist of DNA, a coat (coat protein) for protection, and a means of attachment (terminal filaments). Like other viruses, phages are basically nothing more than packaged DNA. One or more bacteriophages attach to a receptor on the surface of the outer cell membrane of a bacterium. The figure shows how an attached phage inserts its DNA into a bacterium. Numerous different phages are known,e.g., for Escherichia coli and Salmonella (phages T1, T2, P1, F1, lambda, T4, T7, phiX174 and others).

B. Lytic and lysogenic cycles of a bacteriophage

Phages do not reproduce by cell division like bacteria, but by intracellular formation and assembly of the different components. This begins with the attachment of a phage particle to a specific receptor on the surface of a sensitive bacterium. Different phages use different receptors, thus giving rise to specificity of interaction (restriction). The invading phage DNA contains the information for production of coat proteins for new phages and factors for DNA replication and transcription. Translation is provided for by cell enzymes. The phage DNA and phage protein synthesized in the cell are assembled into new phage particles. Finally, the cell disintegrates (lysis) and hundreds of phage particles are released. With attachment of a new phage to a new cell, the procedure is repeated (lytic cycle). Phage reproduction does not always occur after invasion of the cell. Occasionally, phage DNA is integrated into the bacterial chromosome and replicated with it (lysogenic cycle). Phage DNA that has been integrated into the bacterial chromosome is designated a prophage. Bacteria containing prophages are designated lysogenic bacteria; the corresponding phages are termed lysogenic phages. The change from a lysogenic to a lytic cycle is rare. It requires induction by external influences and complex genetic mechanisms.

C. Insertion of a lambda phage into the bacterial chromosome by crossing-over

A phage can be inserted into a bacterial chromosome by different mechanisms. With the lambda phage ( λ ), insertion results from crossing-over between the E. coli chromosome and the lambda chromosome. First, the lambda chromosome forms a ring. Then it attaches to a homologous section of the bacterial chromosome. Both the bacterial and the lambda chromosome are opened by a break and attach to each other. Since the homologies between the two chromosomes are limited to very small regions, phage DNA is seldom integrated. The phage is released (and the lytic cycle is induced) by there verse procedure.(Figures adapted from Watson et al., 1987).