Intercellular Symplastic Connection and Isolation of the Unloading Zone in Flesh of the Developing Grape Berry

The ultrastructure and intercellular connection of the sugar unloading zone (i.e. the phloem in the dorsal vascular bundle and the phloem_surrounding the assimilate sink_cells) of grape ( Vitis vinifera× V. labrusca cv. Jingchao) berry was observed via transmission electron microscopy. The results showed that during the early developmental stages of grape berry, numerous plasmodesmata were found in the phloem between sieve element (SE) and companion cell (CC), between SE/CC complexes, between SE/CC complex and phloem parenchyma cell and in between phloem parenchyma cells, which made the phloem a symplastic integration, facilitating sugar unloading from sieve elements into both companion cells and phloem parenchyma cells via a symplastic pathway. On the contrary, there was almost no plasmodesma between phloem and its surrounding flesh photoassimilate sink_cells, neither in between the flesh photoassimilate sink_cells giving rise to a symplastic isolation both between phloem and its surrounding flesh photoassimilate sink_cells, as well as among the flesh photoassimilate sink_cells. This indicated that both the sugar unloading from phloem and postphloem transport of sugars should be mainly via an apoplastic pathway. During the ripening stage, most of the plasmodesmata between SE/CC complex and the surrounding phloem parenchyma cells were shown to be blocked by the electron_opaque globules, and a phenomenon of plasmolysis was found in a number of companion cells, indicating a symplastic isolation between SE/CC complex and its surrounding parenchyma cells during this phase. The symplastic isolation between the whole phloem and its surrounding photoassimilate sink_cells during the early developmental stages shifted to a symplastic isolation within the phloem during the ripening phase, and thus the symplastic pathway of sugar unloading from SE/CC complex during the early development stages should be replaced by a dominant apoplastic unloading pathway from SE/CC complex in concordance.