Identification of candidate virulence loci in Striga hermonthica, a devastating parasite of African cereal crops
Parasites have evolved suites of proteins, Virulence Factors (VFs), that are delivered into host plants to facilitate colonization. Whilst VFs mediating plant-microbe and plant-nematode interactions have been characterised extensively, less is known about VFs mediating parasitic plant interactions with their hosts. Striga hermonthica is an obligate, root-parasitic plant capable of parasitizing multiple cereal hosts in sub-Saharan Africa, causing devastating losses in yields. An understanding of the molecular nature and allelic variation of VFs in S. hermonthica is essential for breeding durable resistance and delaying the evolution of parasite virulence. To address this issue, we assembled a genome for Striga hermonthica and identified candidate VFs by combining in silico prediction of secreted proteins with pooled sequencing of parasites growing on a susceptible and a strongly resistant rice host. Consistent with predictions for parasites, like S. hermonthica, that can interact with multiple hosts, we identified multiple loci, potentially with a wide range of functions, implicated in overcoming host resistance. Extremely different allele frequencies were observed at 152 non-secreted and 38 putatively secreted VFs between S. hermonthica parasitising the resistant and susceptible rice varieties. Our candidate, secreted VFs encompassed functions such as host cell wall modification, protease inhibitors, oxidoreductase and kinase activities, as well as several with unknown functions. Consistent with maintenance of variation at virulence loci by balancing selection the candidate loci had significantly higher Tajima’s D on average than the genomic background. Our results show that diverse strategies are used by S. hermonthica to overcome different layers of host resistance. Understanding the maintenance of variation at virulence loci by balancing selection will be critical to managing the evolution of virulence as a part of a sustainable control strategy.