Role of starch in carbon translocation and partitioning at the plant level

Endogenous regulation of translocation and of carbon partitioning, major factors for integrating plant function, depend on diurnal regulation of starch synthesis and mobilization. Regulated diurnal cycling of transitory starch provides a steady carbon supply to sink growth and avoids potentially adverse high sugar levels. Carbon availability from starch affects development and alters carbon availability with respect to nitrogen. Along with sugar sensing, the level and turnover of starch are involved in endogenous regulation in response to carbohydrate status. Despite their key roles in plant metabolism, mechanisms for endogenous regulation of starch synthesis and degradation are not well characterized. Time course studies with labeled carbon reveal endogenous diurnal regulation of starch metabolism, by which sucrose synthesis from starch and newly-fixed carbon are mutually regulated in support of translocation at night, under low light, and during periods of water stress. Even under steady irradiance, which supports photosynthesis at midday levels, starch synthesis begins gradually and slows under an end-of-day circadian regulation that anticipates the dark period. Studies with Arabidopsis mutants identified two requisite components of starch mobilization, endoamylase, and glucose transport across the chloroplast inner envelope. Time course studies of carbohydrate levels and labeling studies of plant-level carbon metabolism in mutant plants with impaired ability to mobilize starch identified steps in starch mobilization that support diurnal regulation of translocation. Endogenously regulated exit of glucose across the chloroplast membrane appears to regulate starch mobilization.