5.6.1  Introduction

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Photoassimilate removal from phloem and delivery to recipient sink cells (phloem unloading) is the final step in photoassimilate transport from source to sink. Within sink cells, cellular metabolism and compartmentation are the end-users of phloem-imported photoassimilates. Combined activities of these sink-located transport and transfer events determine the pattern of photoassimilate partitioning between competing sinks and hence contribute to crop yield (Sections 5.3 and 5.4).

Phloem unloading describes transport events responsible for assimilate movement from se–cc complexes to recipient sink cells. A distinction must be made between transport across the se–cc complex boundary and subsequent movement to recipient sink cells. The former transport event is termed sieve element unloading and the latter post-sieve element transport. On reaching the cytoplasm of recipient sink cells, imported photoassimilates can enter metabolic pathways or be compartmented into organelles (e.g. amyloplasts, protein bodies and vacuoles). Metabolic fates for photoassimilates include catabolism in respiratory pathways, biosynthesis (maintenance and growth) and storage as macromolecules (starch and fructans).

Compared with phloem loading, phloem unloading and subsequent sink utilisation of imported photoassimilates operate within a much broader range of configurations:

1. morphological (e.g. apices, stems, roots, vegetative storage organs, reproductive structures);

2. anatomical (e.g. provascular differentiating sieve elements, protophloem sieve elements lacking companion cells, metaphloem se–cc complexes);

3. developmental (e.g. cell division, cell expansion);

4. metabolic (e.g. storage of soluble compounds/polymers, growth sinks).

A correspondingly large range of strategies for phloem unloading and sink utilisation must be anticipated.

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