Like many living organisms, plants have tissues that specialize in specific plant functions. For example, shoots are well adapted to rapid growth, while roots are structured to acquire resources efficiently. However, sugar production is primarily restricted to the photosynthetic cells that reside in the leaves of angiosperm plants. Sugar and other resources are transported from photosynthetic tissues to other specialized tissues by a process called translocation.
Within a plant, tissues that produce more sugar than they consume are sugar sources – leaves are the primary example of this. Roots, shoots, flowers, and fruits are usually considered to be sugar sinks, as they require more sugar than they can make. Translocation distributes sugar, hormones, amino acids, and some signaling molecules from sugar sources to sugar sinks through a tube-like structure of vascular plants called phloem. Flow can be bidirectional in the phloem, which is composed of cells joined end-to-end by plasmodesmata to form the sieve-tube elements. These cells have thickened cell walls, giving them mechanical support, and are accompanied by neighboring companion cells that facilitate phloem health and loading of solutions into the phloem from surrounding tissues.
Phloem loading can occur via the apoplastic or symplastic routes and may be either passive or active. These pathways to phloem may operate at the same time or sequentially, and there is some evidence that plants can switch between loading modes depending on plant water and energy demands. In many instances, the sucrose/H+ symporter couples the loading of sucrose into the phloem with transport of a hydrogen ion.
According to the pressure-flow hypothesis, the sugar concentration gradient promotes the flow of water into the phloem, resulting in the generation of pressure. As a result, the phloem sap moves towards areas of lower pressure, at the nearest sugar sink. At the sugar sink, sucrose is transported to the area of lower sugar concentration, driving movement out of the phloem. Water follows the sucrose, relieving pressure in the phloem.
