P1 phage

P1 is a temperate bacteriophage that infects Escherichia coli and some other bacteria. When undergoing a lysogenic cycle the phage genome exists as a plasmid in the bacterium unlike other phages (e.g. the lambda phage) that integrate into the host DNA. P1 has an icosahedral head containing the DNA attached to a contractile tail with six tail fibers. The P1 phage has gained research interest because it can be used to transfer DNA from one bacterial cell to another in a process known as transduction. As it replicates during its lytic cycle it captures fragments of the host chromosome. If the resulting viral particles are used to infect a different host the captured DNA fragments can be integrated into the new host's genome. This method of in vivo genetic engineering was widely used for many years and is still used today, though to a lesser extent. P1 can also be used to create the P1-derived artificial chromosome cloning vector which can carry relatively large fragments of DNA. P1 encodes Cre recombinase, which is widely used to carry out cell-specific or time-specific DNA recombination by flanking the target DNA with loxP sites.

The virion is similar in structure to the T4 phage but simpler. It has an icosahedral head containing the genome attached at one vertex to the tail. The tail has a tube surrounded by a contractile sheath. It ends in a base plate with six tail fibres. The tail fibres are involved in attaching to the host and providing specificity.

The genome of the P1 phage is moderately large, around 93kbp in length (compared to the genomes of e.g. T4 – 169kbp, lambda – 48kbp and Ff – 6.4kbp). In the viral particle it is in the form of a linear double stranded DNA molecule. Once inserted into the host it circularizes and replicates as a plasmid.

In the viral particle the DNA molecule is longer (110kbp) than the actual length of the genome. It is created by cutting an appropriately sized fragment from a concatemeric DNA chain having multiple copies of the genome (see the section below on lysis for how this is made). Due to this the ends of the DNA molecule are identical. This is referred to as being terminally redundant. This is important for the DNA to be circularized in the host. Another consequence of the DNA being cut out of a concatemer is that a given linear molecule can start at any location on the circular genome. This is called a cyclic permutation.

The genome is especially rich in Chi sequences recognized by the bacterial recombinase RecBCD. The genome contains two origins of replication: oriR which replicates it during the lysogenic cycle and oriL which replicates it during the lytic stage. The genome of P1 encodes three tRNAs which are expressed in the lytic stage.

The genome of P1 encodes 112 proteins and 5 untranslated genes and is this about twice the size of bacteriophage lambda.

The phage particle adsorbs onto the surface of the bacterium using the tail fibers for specificity. The tail sheath contracts and the DNA of the phage is injected into the host cell. The host DNA recombination machinery or the cre enzyme translated from the viral DNA recombine the terminally redundant ends and circularize the genome. Depending on various physiological cues, the phage may immediately proceed to the lytic phase or it may enter a lysogenic state.

The gene that encodes the tail fibers have a set of sequences that can be targeted by a site specific recombinase Cin. This causes the C terminal end of the protein to switch between two alternate forms at a low frequency. The viral tail fibers are responsible for the specificity of binding to the host receptor. The targets of the viral tail fibers are under a constant pressure to evolve and evade binding. This method of recombinational diversity of the tail allows the virus to keep up with the bacterium. This system has close sequence homologies to recombinational systems in the tail fibers of unrelated phages like the mu phage and the lambda phage.