, 2005). This number corresponds to a small minority of the S. cerevisiae genome (< 1%); however, these genes have contributed to important functional innovations, including the ability to synthesize biotin, the ability to grow under anaerobic conditions and the ability to utilize Lumacaftor research buy sulphate from several organic sources (Hall et al., 2005). Similarly, a recent sequencing project of the commercial wine yeast strain EC118 uncovered three genomic regions that have been transferred horizontally from other

fungal sources (Novo et al., 2009). The three regions encode 34 genes, which are important in wine fermentation including nitrogen and carbon metabolism, cellular transport and stress responses, that aid yeast wine strains adapt to high sugar, low nitrogen and high ethanol concentrations (Novo et al., 2009). Other HGT events that have contributed to niche specification include the acquisition of glycosyl hydrolases (GHs) by rumen fungi from prokaryotes (Garcia-Vallve et al., 2000). GHs have permitted rumen fungi to establish a niche in the rumen of herbivorous mammals where cellulose and plant hemicellulose are

the main carbon sources (Garcia-Vallve et al., 2000). Similarly, the entomopathogenic fungus Metarhizium anisopliae has acquired a phosphoketolase (Mpk1) from an unspecified Epigenetics inhibitor bacterial source. It has been demonstrated that Mpk1 is necessary for insect virulence and is highly expressed in trehalose-rich insect haemolymph, thus playing an important role in niche adaptation for this fungus in the insect haemocoel. Slot & Hibbett (2007) have also uncovered an ancient transfer of a nitrate assimilation cluster from the Oomycota to an ancestral Dikarya species and propose that the acquisition of high-affinity nitrate assimilation contributed to the success of Dikarya on land by allowing exploitation Erastin nmr of nitrate in aerobic soils. Furthermore, the subsequent transfer of a complete Basidiomycete nitrate assimilation cluster into the ascomycetous mould Trichoderma reesei improved fitness and corresponds to a change in nutritional mode (wood decayer), providing

further evidence that horizontal transfer can facilitate niche shift in fungi (Slot & Hibbett, 2007). Incidences of HGT have also been linked to virulence in fungi, and the recent acquisition of a toxin gene (ToxA) by Pyrenophora tritici-repentis from Stagonospora nodorum has resulted in serious Pyrenophora infestations of wheat (Friesen et al., 2006). ToxA exerts its toxic effect via internalization into sensitive wheat mesophyll cells where it localizes to chloroplasts (Manning & Ciuffetti, 2005); however, the mechanisms involved in ToxA-mediated cell death remain to be elucidated. Interfungal HGT of a pea pathogenicity gene (PEP) cluster from Fusarium oxysporum to Nectria haematococca has also been linked to disease. The PEP cluster increases pathogenicity by converting a pea phytoalexin (pisatin) into a less toxic compound (Matthews & Van Etten, 1983).