Inorganic phosphate (Pi) deficiency is a major limiting factor for plant growth and development. Previous reports have demonstrated that PHOSPHATE STARVATION RESPONSE 1 (PHR1) and OsPHR2 play central roles in Pi-starvation signaling in Arabidopsis and rice, respectively. However, the Pi-starvation signaling network in tomato (Solanum lycopersicum) is still not fully understood. In this work, SlPHL1, a homolog of AtPHR1 and OsPHR2, was identified from tomato. It was found that SlPHL1 contains the MYB and coiled-coil (CC) domains, localizes in the nucleus, and has transcriptional activity, indicating that it is a typical MYB-CC transcription factor (TF). Overexpression of SlPHL1 enhanced Pi-starvation responses both in Arabidopsis Col-0 and in tomato Micro-Tom, including elevated root hair growth, promoted APase activity, favored Pi uptake, and increased transcription of Pi starvationinducing (PSI) genes. Besides, overexpressing SlPHL1 was able to compensate for the Pi-starvation response weakened by the AtPHR1 mutation. Notably, electrophoretic mobility shift assay (EMSA) showed that SlPHL1 could bind to the PHR1-binding sequence (P1BS, GNATATNC)-containing DNA fragments. Furthermore, SlPHL1 specifically interacted with the promoters of the tomato PSI genes SlPht1;2 and SlPht1;8 through the P1BS cis-elements. Taken these results together, SlPHL1 is a newly identified MYB-CC TF from tomato, which participates in Pi-starvation signaling by directly upregulating the PSI genes. These findings might contribute to the understanding of the Pi-starvation signaling in tomato. | INTRODUCTIONAs an essential macronutrient, phosphorus (P) is a key component in numerous molecules, including nucleic acids, proteins, phospholipids, ATP, and NADPH, and it participates in plant energy metabolism, photosynthesis, and respiration (Wang et al., 2018). P in soil mainly exists in two forms, namely inorganic phosphate (Pi, mainly H 2 PO 4 À and HPO 4 2À

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Introduction Diabetic kidney disease (DKD) is a major microvascular complication of diabetes mellitus (DM) and the primary cause of end-stage renal disease. Routine medication might result in adverse effects which limit their usage. Laboratory studies have shown that Xiao Zheng San Jie (XZSJ) granules is effective in reducing the levels of urinary protein and blood creatinine in rats with DKD. Hence, we aim to conduct a clinical trial to determine the efficacy and safety of XZSJ granules among patients with DKD Stage IV, who present with the syndromes of qi deficiency and blood stasis. The purpose of this paper is to report the methodology of this trial.Methods and analysisThis is a randomised, double-blind, placebo-controlled, multi-centre clinical trial. A total of 72 patients with DKD Stage IV who have the syndromes of qi deficiency and blood stasis will be randomly assigned to either intervention or placebo group. In addition to routine medication, the intervention group will be given XZSJ formula (in granules) and the control group will be given placebo for 12-weeks. Data collected at baseline as well as weeks 4, 8 and 12 will be recorded in case report forms and subsequently entered into EpiData 3.1 electronic database. Adverse events will also be documented. Primary outcomes – estimated glomerular filtration rate, urinary albumin-to-creatinine ratio, and serum creatinine – will be assessed after 12 weeks. Ethics and disseminationThis trial has been approved by the Research Ethics Committee of Dongzhimen Hospital (DZMEC-KY-2018-60). Informed consent will be obtained from all participants. Results of this study will be disseminated to the public through academic conferences and peer-reviewed journals.DiscussionThis trial might provide information on an effective method for the treatment of DKD, especially for patients who present with the syndromes of qi deficiency and blood stasis. We hope that this trial will serve as a reference for the conception of high-quality Traditional Chinese Medicine trials in the future, which incorporate syndrome diagnosis into the study design.Trial registration Chinese Clinical Trial Registry, ChiCTR1900021391. Registered 18 February 2018, http://www.chictr.org.cn/showproj.aspx?proj=36106.