envisaged to be contributed to by variation in selection, mutational bias, and biased recombination-associated DNA repair. The average GC-content in human genomes ranges from 35% to 60% across 100-Kb fragments, with a mean of 41%. The GC-content of Yeast (Saccharomyces cerevisiae) is 38%, and that of another common model organism, thale cress (Arabidopsis thaliana), is 36%. Because of the nature of the genetic code, it is virtually impossible for an organism to have a genome with a GC-content approaching either 0% or 100%. However, a species with an extremely low GC-content is Plasmodium falciparum (GC% = ~20%), and it is usually common to refer to such examples as being AT-rich instead of GC-poor. Several mammalian species (e.g., shrew, microbat, tenrec, rabbit) have independently undergone a marked increase in the GC-content of their genes. These GC-content changes are correlated with species life-history traits (e.g., body mass or longevity) and genome size, and might be linked to a molecular phenomenon called the GC-biased gene conversion. Applications Molecular biology In polymerase chain reaction (PCR) experiments, the GC-content of short oligonucleotides known as primers is often used to predict their annealing temperature to the template DNA. A higher GC-content level indicates a relatively higher melting temperature. Many sequencing technologies, such as Illumina sequencing, have trouble reading high-GC-content sequences. Bird genomes are known to have many such parts, causing the problem of "missing genes" expected to be present from evolution and phenotype but never sequenced — until improved methods were used. Systematics The species problem in non-eukaryotic taxonomy has led to various suggestions in classifying bacteria, and the ad hoc committee on reconciliation of approaches to bacterial systematics of 1987 has recommended use of GC-ratios in higher-level hierarchical classification. For example, the Actinomycetota are characterised as "high GC-content bacteria". In Streptomyces coelicolor A3(2), GC-content is 72%. With the use of more reliable, modern methods of molecular systematics, the GC-content definition of Actinomycetota has been abolished and low-GC bacteria of this clade have been found. Software tools GCSpeciesSorter and TopSort are software tools for classifying species based on their GC-contents. See also Codon usage bias References External links Table with GC-content of all sequenced prokaryotes Taxonomic browser of bacteria based on GC ratio on NCBI website. GC ratio in diverse species. DNA Molecular biology Biological classification