Sucrose transporters and plasmodesmal regulation in passive phloem loading

An essential step for the distribution of carbon throughout the whole plant is the loading of sugars into the phloem in source organs. In many plants, accumulation of sugars in the sieve element-companion cell (SE-CC) complex is mediated and regulated by active processes. However, for poplar and many other tree species, a passive symplasmic mechanism of phloem loading has been proposed, characterized by symplasmic continuity along the pre-phloem pathway and the absence of active sugar accumulation in the SE-CC complex. A high overall leaf sugar concentration is thought to enable diffusion of sucrose into the phloem. In this review, we critically evaluate current evidence regarding the mechanism of passive symplasmic phloem loading, with a focus on the potential influence of active sugar transport and plasmodesmal regulation. The limited experimental data, combined with theoretical considerations, suggest that a concomitant operation of passive symplasmic and active phloem loading in the same minor vein is unlikely. However, active sugar transport could well play an important role in how passively loading plants might modulate the rate of sugar export from leaves. Insights into the operation of this mechanism has direct implications for our understanding of how these plants utilize assimilated carbon.