Phloem unloading follows an extensive apoplastic pathway in developing strawberry fruit

Strawberry (Fragaria X ananassa) is widely used as a model for fleshy fruit biology research; nevertheless, the pathway of phloem unloading in strawberry fruit remains unknown. Using transmission electron microscopy (TEM) and the symplastic tracer, carboxyfluorescein, the phloem unloading pathway was studied in 'Akihime' strawberry fruit from post-anthesis to the ripening stage. Ultrastructural analysis showed that many plasmodesmata existed between the sieve element cells (SEC) and the companion cells (CC), but plasmodesmata were absent between the sieve element cell - CC complexes and their surrounding phloem parenchyma cells (PPCs). Interestingly, an abundance of plasmodesmata was observed between parenchyma cells. Imaging of carboxyfluorescein unloading revealed that the dye was confined only to the phloem vascular bundles in developing receptacles. In conclusion, the results obtained from the present study indicate that sugar unloading from the phloem occurs through an apoplastic pathway. Post-phloem transport of sugar then follows a symplastic pathway in strawberry fruit.