What Is Phloem Unloading?

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Abstract
Several studies of phloem unloading have failed to distinguish between transport events occurring at the sieve element/companion cell boundary and subsequent short-distance transport through parenchyma cells. Indirect evidence has been obtained for symplastic unloading in storage and utilization sinks. In other sinks transfer to the apoplast may occur, but not necessarily at the sieve element/companion cell complex, and the evidence for apoplastic phloem unloading is equivocal, as is the role of apoplastic acid invertase in this process. The ability of several types of sink cells to accumulate sugars from the apoplast is discussed in the conflicting light of functional symplastic continuity between sink cells. Attention is drawn to the complexity of the postunloading pathway in many sinks and the difficulty of determining the exact sites of symplast/apoplast solute exchange. Potential future areas for study in the field are highlighted.