In the evolution of land plants, the plant immune system has experienced expansion in immune receptor and signaling pathways. Lineage-specific expansions have been observed in diverse gene families that are potentially involved in immunity but lack causal association. Here, we show that Rps8-mediated resistance in barley to the pathogen Puccinia striiformis f. sp. tritici (wheat stripe rust) is conferred by a genetic module: Pur1 and Exo70FX12, which are together necessary and sufficient. Pur1 encodes a leucine-rich repeat receptor kinase and is the ortholog of rice Xa21, and Exo70FX12 belongs to the Poales-specific Exo70FX clade. The Exo70FX clade emerged after the divergence of the Bromeliaceae and Poaceae and comprises from 2 to 75 members in sequenced grasses. These results demonstrate the requirement of a lineage-specific Exo70FX12 in Pur1-mediated immunity and suggest that the Exo70FX clade may have evolved a specialized role in receptor kinase signaling.

Australian Tropical Herbarium James Cook University Smithfield 4878 Australia

Centre of the Region Haná for Biotechnological and Agricultural Research Institute of Experimental Botany of the Czech Academy of Sciences Šlechtitelů 31 779 00 Olomouc Czech Republic

Department of Agricultural and Food Sciences University of Bologna Viale G Fanin 44 40127 Bologna Italy

Department of Plant Physiology RWTH Aachen University 52056 Aachen Germany

NIAB 93 Lawrence Weaver Road Cambridge CB3 0LE England UK

The Sainsbury Laboratory University of East Anglia Norwich Research Park Norwich NR4 7UH UK

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