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Cereal cyst nematodes are sedentary biotrophic endoparasites that secrete effector proteins into plant tissues to transit normal cells into specialized feeding sites and suppress plant defenses. To understand the function of nematode effectors in  Heterodera avenae , here, we identified a calreticulin protein HaCRT1, which could suppress the cell death induced by Bax when expressed in  Nicotiana benthamiana . HaCRT1 is synthetized in the subventral gland cells of pre-parasitic second-stage nematodes. Real-time PCR assays indicated that the expression of  HaCRT1  was highest in parasitic second-stage juveniles. The expression of an HaCRT1-RFP fusion in  N. benthamiana  revealed that it was localized in the endoplasmic reticulum of the plant cell. The ability of  H. avenae  infecting plants was significantly reduced when  HaCRT1  was knocked down by RNA interference  in vitro .  Arabidopsis thaliana  plants expressing  HaCRT1  were more susceptible than wild-type plants to  Pseudomonas syringae . The induction of defense-related genes,  PAD4 ,  WRKY33 ,  FRK1 , and  WRKY29 , after treatment with flg22 was suppressed in  HaCRT1 -transgenic plants. Also, the ROS accumulation induced by flg22 was reduced in the  HaCRT1 -transgenic plants compared to wild-type plants. HaCRT1 overexpression increased the cytosolic Ca 2+  concentration in  A. thaliana . These data suggested that HaCRT1 may contribute to the pathogenicity of  H. avenae  by suppressing host basal defense.

HaCRT1 of Heterodera avenae Is Required for the Pathogenicity of the Cereal Cyst Nematode