Plastidic metabolite transporters and their physiological functions in theinducible crassulacean acid metabolism plant Mesembryanthemum crystallinum

The inducible crassulacean acid metabolism (CAM) plant Mesembryanthemum cry stallinum accumulates malic acid during the night and converts it to starch during the day via a pathway that, because it is located in different subc ellular compartments, depends on specific metabolite transport across membr anes. The chloroplast glucose transporter (pGlcT) and three members of the phosphate translocator (PT) family were isolated. After induction of CAM, t ranscript amounts of the phosphoenolpyruvate (PEP) phosphate translocator ( PPT) and the glucose-g-phosphate (Glc6P) phosphate translocator (GPT) genes were increased drastically, while triose phosphate (TP) phosphate transloc ator (TPT) and the pGlcT transcripts remained unchanged. PPT- and GPT-speci fic transcripts and transporter activities exhibited a pronounced diurnal v ariation, displaying the highest amplitude in the light. pGlcT transcripts were elevated towards the end of the light period and at the beginning of t he dark period. These findings, combined with diurnal variations of enzyme activities and metabolite contents, helped to elucidate the roles of the PP T, GPT, TPT and pGlcT in CAM. The main function of the PPT is the daytime e xport from the stroma of PEP generated by pyruvate orthophosphate:dikinase (PPDK). The increased transport activity of GPT in the light suggests a hig her requirement for Glc6P import for starch synthesis rather than starch mo bilization. Most likely, Glc6P rather than 3-phosphoglycerate or triose pho sphates is the main substrate for daytime starch biosynthesis in M. crystal linum plants in which CAM has been induced (CAM-induced), similar to non-gr een plastids. In the dark, starch is mobilized both phosphorylytically and amylolytically and the products are exported by the GPT, TPT and pGlcT. The transport activities of all three phosphate translocators and the transcri pt amounts of the pGlcT adapt to changing transport requirements in order t o maintain high metabolic fluxes during the diurnal CAM cycle.