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The earliest proteolytic event affecting most proteins is the excision of the initiating Met (NME). This is an essential and ubiquitous cotranslational process tightly regulated in all eukaryotes. Currently, the effects of NME on unknown complex cellular networks and the ways in which its inhibition leads to developmental defects and cell growth arrest remain poorly understood. Here, we provide insight into the earliest molecular mechanisms associated with the inhibition of the NME process in Arabidopsis thaliana. We demonstrate that the developmental defects induced by NME inhibition are caused by an increase in cellular proteolytic activity, primarily induced by an increase in the number of proteins targeted for rapid degradation. This deregulation drives, through the increase of the free amino acids pool, a perturbation of the glutathione homeostasis, which corresponds to the earliest limiting, reversible step promoting the phenotype. We demonstrate that these effects are universally conserved and that the reestablishment of the appropriate glutathione status restores growth and proper development in various organisms. Finally, we describe a novel integrated model in which NME, protein N-alpha-acylation, proteolysis, and glutathione homeostasis operate in a sequentially regulated mechanism that directs both growth and development.

We analyzed the physiological impact of function loss on cheesmaniae alleles at the HKT1;1 and HKT1;2 loci in the roots and aerial parts of tomato plants in order to determine the relative contributions of each locus in the different tissues to plant Na+/K+ homeostasis and subsequently to tomato salt tolerance. We generated different reciprocal rootstock/scion combinations with non-silenced, single RNAi-silenced lines for ScHKT1;1 and ScHKT1;2 , as well as a silenced line at both loci from a near isogenic line (NIL14), homozygous for the Solanum cheesmaniae haplotype containing both HKT1 loci and subjected to salinity under natural greenhouse conditions. Our results show that salt treatment reduced vegetative growth and altered the Na+/K+ ratio in leaves and flowers; negatively affecting fruit production, particularly in graft combinations containing single silenced ScHKT1;2 - and double silenced S c HKT1; 1 /ScHKT1; 2 lines when used as scion. We concluded that the removal of Na+ from the xylem by ScHKT1;2 in the aerial part of the plant can have an even greater impact than that on Na+ homeostasis at the root level under saline conditions. Also, ScHKT1;1 function loss in rootstock greatly reduced the Na+/K+ ratio in leaf and flower tissues, minimized yield loss under salinity. Our results suggest that, in addition to xylem Na+ unloading, ScHKT1;2 could also be involved in Na+ uploading into the phloem, thus promoting Na+ recirculation from aerial parts to the roots. This recirculation of Na+ to the roots through the phloem could be further favoured by ScHKT1;1 silencing at these roots. • Physiological impact of ScHKT1;1/ScHKT1;2 function loss in roots and aerial parts on tomato salt tolerance in a commercial greenhouse. • Reciprocal grafting combinations with a NIL, homozygous for ScHKT1;1 / ScHKT1;2 and their respective single and double RNAi lines. • ScHKT1;2 alleviates the negative saline effects on growth and fruit yields. • Main function of ScHKT1;2 is xylem Na+ removal at the aerial part. • ScHKT1;1 function loss in rootstock sharply reduced the Na+/K+ ratio in leaves and flowers. [ABSTRACT FROM AUTHOR]