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Open Access Highly Accessed Research article

Motif depletion in bacteriophages infecting hosts with CRISPR systems

Anne Kupczok12* and Jonathan P Bollback1

Author Affiliations

1 IST Austria, Am Campus 1, 3400 Klosterneuburg, Austria

2 Institute of Microbiology, Christian-Albrechts-University of Kiel, 24118 Kiel, Germany

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BMC Genomics 2014, 15:663  doi:10.1186/1471-2164-15-663

Published: 8 August 2014



CRISPR is a microbial immune system likely to be involved in host-parasite coevolution. It functions using target sequences encoded by the bacterial genome, which interfere with invading nucleic acids using a homology-dependent system. The system also requires protospacer associated motifs (PAMs), short motifs close to the target sequence that are required for interference in CRISPR types I and II. Here, we investigate whether PAMs are depleted in phage genomes due to selection pressure to escape recognition.


To this end, we analyzed two data sets. Phages infecting all bacterial hosts were analyzed first, followed by a detailed analysis of phages infecting the genus Streptococcus, where PAMs are best understood. We use two different measures of motif underrepresentation that control for codon bias and the frequency of submotifs. We compare phages infecting species with a particular CRISPR type to those infecting species without that type. Since only known PAMs were investigated, the analysis is restricted to CRISPR types I-C and I-E and in Streptococcus to types I-C and II. We found evidence for PAM depletion in Streptococcus phages infecting hosts with CRISPR type I-C, in Vibrio phages infecting hosts with CRISPR type I-E and in Streptococcus thermopilus phages infecting hosts with type II-A, known as CRISPR3.


The observed motif depletion in phages with hosts having CRISPR can be attributed to selection rather than to mutational bias, as mutational bias should affect the phages of all hosts. This observation implies that the CRISPR system has been efficient in the groups discussed here.

Bacterial immunity; Bacteria-phage coevolution; Selection; PAM