Phloem: the long and the short of it

The structure of phloem sieve-element-companion-cell complexes reflects a duality of function: to conduct photoassimilates throughout the plant, and to exchange solutes between the phloem and surrounding tissues. The conceptual integration of these long-and short-distance functions requires the abandonment of a long-cherished concept in phloem physicochemistry, that source-sink turgor differentials control flow. The manifest inability of decentralized organisms such as plants to control phloem translocation centrally disqualifies such differentials as control variables; besides, the phloem is maximally efficient if the pressure differentials are small. Testing this hypothesis and whether turgor differentials are small will require a significant recommitment to studying the quantitative anatomy of phloem.