The central circadian regulator CCA1 functions in Glycine max during defense to a root pathogen, regulating the expression of genes acting in effector triggered immunity (ETI) and cell wall metabolism

Abstract

Expression of the central circadian oscillator components CIRCADIAN CLOCK ASSOCIATED 1 (CCA1), TIMING OF CAB1 (TOC1), GIGANTEA (GI), and CONSTANS (CO) occurs in Glycine max root cells (syncytia) parasitized by the nematode Heterodera glycines while undergoing resistance, indicating a defense role. GmCCA1-1 relative transcript abundance (RTA) in roots experiencing overexpression (OE) or RNA interference (RNAi) is characterized by rhythmic oscillations, compared to a ribosomal protein gene (GmRPS21) control. A GmCCA1-1 RTA change, advancing by 12 h, exists in H. glycines-infected as compared to uninfected controls in wild-type, H. glycines-resistant, G. max_{[Peking/PI 548402]}. The G. max_{[Peking/PI 548402]} transgenic controls exhibit the RTA change by 4 h post infection (hpi), not consistently occurring in the H. glycines-susceptible G. max_{[Williams 82/PI 518671]} until 56 hpi. GmCCA1-1 expression is observed to be reduced in H. glycines-infected GmCCA1-1-OE roots as compared to non-infected transgenic roots with no significant change observed among RNAi roots. The GmCCA1-1 expression in transgenic GmCCA1-1-OE roots remains higher than control and RNAi roots. Decreased GmCCA1-1 mRNA among infected roots shows the altered expression is targeted by H. glycines. Gene expression of proven defense genes including 9 different mitogen activated protein kinases (GmMAPKs), NON-RACE SPECIFIC DISEASE RESISTANCE 1 (GmNDR1-1), RPM1-INTERACTING PROTEIN 4 (GmRIN4-4), and the secreted xyloglucan endotransglycosylase/hydrolase 43 (GmXTH43) in GmCCA1-1-OE and GmCCA1-1-RNAi roots, compared to controls, reveal a significant role of GmCCA1-1 expression in roots undergoing defense to H. glycines parasitism. The observation that GmCCA1-1 regulates GmXTH43 expression links the central circadian oscillator to the functionality of the secretion system.

Keywords: CIRCADIAN CLOCK ASSOCIATED 1 (CCA1); CONSTANS (CO); Central circadian oscillator; Clock; Defense; GIGANTEA (GI); Heterodera glycines; Mitogen activated protein kinase (MAPK); NON-RACE SPECIFIC DISEASE RESISTANCE 1 (NDR1); Nematode; Root; Soybean; TIMING OF CAB1 (TOC1); Xyloglucan endo-transglycosylase/hydrolase (XTH).