Molecular mechanisms regulating Pi-signaling and Pi homeostasis under OsPHR2,a central Pi-signaling regulator,in rice

Phosphorus(P)is one of the most important major mineral elements for plant growth and metabolism.Plants have evolved adaptive regulatory mechanisms to maintain phosphate(Pi)homeostasis by improving phosphorus uptake,translocation,remobilization and efficiency of use.Here we review recent advances in our understanding of the OsPHR2-mediated phosphate-signaling pathway in rice.OsPHR2 positively regulates the low-affinity Pi transporter OsPT2 through physical interaction and reciprocal regulation of OsPHO2 in roots.OsPT2 is responsible for most of the OsPHR2-mediated accumulation of excess Pi in shoots.OsSPX1 acts as a repressor in the OsPHR2-mediated phosphate-signaling pathway.Some mutants screened from ethyl methanesulfonate(EMS)-mutagenized M2 population of OsPHR2 overexpression transgenic line removed the growth inhibition,indicating that some unknown factors are crucial for Pi utilization or plant growth under the regulation of OsPHR2.

A reciprocal inhibitory module for Pi and iron signaling

Phosphorous(P)and iron(Fe),two essential nutrients for plant growth and development,are highly abundant elements in the earth's crust but often display low availability to plants.Due to the ability to form insoluble complexes,the antagonistic interaction between P and Fe nutrition in plants has been noticed for decades.However,the underlying molecular mechanism modulating the signaling and homeostasis between them re-mains obscure.Here,we show that the possible iron sensors HRZs,the iron deficiency-induced E3 ligases,could interact with the central regulator of phosphate(Pi)signaling,PHR2,and prompt its ubiquitination at lysine residues K319 and K328,leading to its degradation in rice.Consistent with this,the hrzs mutants dis-played a high Pi accumulation phenotype.Furthermore,we found that iron deficiency could attenuate Pi star-vation signaling by inducing the expression of HRZs,which in turn trigger PHR2 protein degradation.Inter-estingly,on the other hand,rice PHRs could negatively regulate the expression of HRZs to modulate iron deficiency responses.Therefore,PHR2 and HRZs form a reciprocal inhibitory module to coordinate Pi and iron signaling and homeostasis in rice.Taken together,our results uncover a molecular link between Pi and iron master regulators,which fine-tunes plant adaptation to Pi and iron availability in rice.

Inositol Pyrophosphate lnsP8 Acts as an Intracellular Phosphate Signal in Arabidopsis

The maintenance of cellular phosphate(Pi)homeostasis is of great importance in living organisms.The SPX domain-containing protein 1(SPX1)proteins from both Arabidopsis and rice have been proposed to act as sensors of Pi status.The molecular signal indicating the cellular Pi status and regulating Pi homeostasis in plants,however,remains to be identified,as Pi itself does not bind to the SPX domain.Here,we report the identification of the inositol pyrophosphate lnsP8 as a signaling molecule that regulates Pi homeostasis in Arabidopsis.Polyacrylamide gel electrophoresis profiling of InsPs revealed that lnsP8 level positively cor­relates with cellular Pi concentration.We demonstrated that the homologs of diphosphoinositol pentaki-sphosphate kinase(PPIP5K),VIH1 and VIH2,function redundantly to synthesize lnsP8,and that the vih1 vih2 double mutant overaccumulates Pi.SPX1 directly interacts with PHR1,the central regulator of Pi star­vation responses,to inhibit its function under Pi-replete conditions.However,this interaction is compro­mised in the vih1 vih2 double mutant,resulting in the constitutive induction of Pi starvation-induced genes,indicating that plant cells cannot sense cellular Pi status without lnsP8.Furthermore,we showed that lnsP8 could directly bind to the SPX domain of SPX1 and is essential for the interaction between SPX1 and PHR1.Collectively,our study suggests that lnsP8 is the intracellular Pi signaling molecule serving as the ligand of SPX1 for controlling Pi homeostasis in plants.

Intraspecific Variations of Phosphorus Absorption and Remobilization, P Forms, and Their Internal Buffering in Brassica Cultivars Exposed to a P-Stressed Environment

Translocation of absorbed phosphorus （P） from metabolically inactive sites to active sites in plants growing under P deprivation may increase its P utilization efficiency （PUE）. Acclimation to phosphate （Pi） starvation may be caused by a differential storage pool of vacuolar P, its release, and the intensity of re-translocation of absorbed P as P starvation inducible environmental cues （PSlEC） from ambient environment. Biomass assay and three P forms, namely inorganic （Pi）, organic （Po）, and acid-soluble total （Ptas） were estimated in Brassica cultivars exposed to 10 d P deprivation in the culture media. Considering that -aPi/at denotes the rate of Pi release, Pi release velocity （RSPi） was determined as the tangent to the equations obtained for Pi f（t） at the mean point in the period of greatest Pi decrease, whereas the inverse of the RSPi was an estimate of the internal Pi buffering capacity （IBCPi）. Inter cultivar variations in size of the non-metabolic Pi pool, RSPi, re-translocation of Pi from less to more active metabolic sites, and preferential Pi source and sink compartments were evaluated under P starvation. The cultivar ＇Brown Raya＇ showed the highest Pi storage ability under adequate external P supply, and a more intensive release than ＇Rain Bow＇ and ＇Dunkled＇ under P stress. Cultivar ＇B.S.A＇ was inferior to ＇Con-l＇ in its ability to store and use Pi. Roots and upper leaves were the main sink of Pi stored in the lower and middle leaves of all cultivars and showed lower IBCPi and larger RSPi values than lower and middle leaves. In another trial, six cultivars were exposed to P-free nutrition for 29 d after initial feeding on optimum nutrition for 15 d. With variable magnitude, all of the cultivars re-translocatad P from the above ground parts to their roots under P starvation, and [P] at 44 d after transplanting was higher in developing leaves compared with developed leaves. Under P deprivation, translocation of absorbed P from metabolically inactive to active sites may have helped the tolerant cultivars to establish a better rooting system, which provided a basis for tolerance against P starvation and increased PUE. A better understanding of the extent to which changes in the flux of P absorption and re-translocation under PSIEC will help to scavenge Pi from bound P reserves and will bring more sparingly soluble P into cropping systems and obtain capitalization of P reserves.