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The small phenolic compound salicylic acid (SA) plays a critical role in plant defense against broad-spectrum of pathogens. The phosphate transporter gene  PHT4;1  was previously shown to affect SA-mediated defense and its expression is regulated by the circadian clock. To further understand how  PHT4;1  affects SA accumulation, here we analyzed the genetic interactions between the gain-of-function mutant  pht4;1-1  and several known SA mutants, including  sid2-1 ,  ald1-1 ,  eds5-3 , and  pad4-1 . The genetic analysis was conducted in the  acd6-1  background since the change of  acd6-1  dwarfism can be used as a convenient readout for the change of defense levels caused by impairments in some SA genes. We found that compared with the corresponding double mutants, the triple mutants  acd6-1pht4;1-1ald1-1 ,  acd6-1pht4;1-1eds5-3 , and  acd6-1pht4;1-1pad4-1  accumulated lower levels of SA and  PR1  transcripts, suggesting that  PHT4;1  contributes to  acd6-1- conferred defense phenotypes independently of these known SA regulators. Although some triple mutants had wild type (wt)-like levels of SA and  PR1  transcripts, these plants were smaller than wt and displayed minor cell death, suggesting that additional regulatory pathways contribute to  acd6-1- conferred dwarfism and cell death. Our data further showed that circadian expression of  PHT4;1  was dependent on CIRCADIAN CLOCK ASSOCIATED 1 (CCA1), a central oscillator component of  Arabidopsis  circadian clock. Recombinant CCA1 protein was demonstrated to bind to the  PHT4;1  promoter in electrophoretic mobility shift assays, suggesting a direct transcriptional regulation of  PHT4;1  by CCA1. Together these results indicate that PHT4;1 is a SA regulator acting independently of several known SA genes and they also implicate a role of the circadian clock mediated by CCA1 in regulating phosphate transport and/or innate immunity in  Arabidopsis .

The phosphate transporter PHT4;1 is a salicylic acid regulator likely controlled by the circadian clock protein CCA1