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Phytophthora root rot, caused by  Phytophthora sojae  is a destructive disease of soybean ( Glycine max ) worldwide. We previously confirmed that the bHLH transcription factor GmPIB1 ( P. sojae -inducible bHLH transcription factor) reduces accumulation of reactive oxygen species (ROS) in cells by inhibiting expression of the peroxidase-related gene  GmSPOD  thus improving the resistance of hairy roots to  P. sojae . To identify proteins interacting with GmPIB1 and assess their participation in the defense response to  P. sojae , we obtained transgenic soybean hairy roots overexpressing GmPIB1 by  Agrobacterium rhizogenes  mediated transformation and examined GmPIB1 protein–protein interactions using immunoprecipitation combined with mass spectrometry. We identified 392 proteins likely interacting with GmPIB1 and selected 20 candidate genes, and only 26S proteasome regulatory subunit GmPSMD (Genbank accession no.  XP_014631720 ) interacted with GmPIB1 in luciferase complementation and pull-down experiments and yeast two-hybrid assays. Overexpression of  GmPSMD  ( GmPSMD- OE) in soybean hairy roots remarkably improved resistance to  P. sojae  and RNA interference of  GmPSMD  ( GmPSMD  -RNAi) increased susceptibility. In addition, accumulation of total ROS and hydrogen peroxide (H 2 O 2 ) in  GmPSMD- OE transgenic soybean hairy roots were remarkably lower than those of the control after  P. sojae  infection. Moreover, in  GmPSMD -RNAi transgenic soybean hairy roots, H 2 O 2  and the accumulation of total ROS exceeded those of the control. There was no obvious difference in superoxide anion (O 2  – ) content between control and transgenic hairy roots. Antioxidant enzymes include peroxidase (POD), glutathione peroxidase (GPX), superoxide dismutase (SOD), catalase (CAT) are responsible for ROS scavenging in soybean. The activities of these antioxidant enzymes were remarkably higher in  GmPSMD -OE transgenic soybean hairy roots than those in control, but were reduced in  GmPSMD -RNAi transgenic soybean hairy roots. Moreover, the activity of 26S proteasome in  GmPSMD -OE and  GmPIB1 -OE transgenic soybean hairy roots was significantly higher than that in control and was significantly lower in  PSMD -RNAi soybean hairy roots after  P. sojae  infection. These data suggest that  GmPSMD  might reduce the production of ROS by improving the activity of antioxidant enzymes such as POD, SOD, GPX, CAT, and  GmPSMD  plays a significant role in the response of soybean to  P. sojae . Our study reveals a valuable mechanism for regulation of the pathogen response by the 26S proteasome in soybean.

The 26S Proteasome Regulatory Subunit GmPSMD Promotes Resistance to Phytophthora sojae in Soybean