The hexose transporters CsHT3 and CsHT16 regulate postphloem transport and fruit development in cucumber

Hexoses are essential for plant growth and fruit development. However, the precise roles of hexose/H+ symporters in postphloem sugar transport and cellular sugar homeostasis in rapidly growing fruits remain elusive. To elucidate the functions of hexose/H+ symporters in cucumber (Cucumis sativus L.) fruits, we conducted comprehensive analyses of their tissue-specific expression, localization, transport characteristics, and physiological functions. Our results demonstrate that CsHT3 (C. sativus hexose transporter), CsHT12, and CsHT16 are the primary hexose/H+ symporters expressed in cucumber fruits. CsHT3 and CsHT16 are localized in the sieve element-companion cell during the ovary and early fruit development stages. As the fruit develops and expands, the expression of both symporters shifts to phloem parenchyma cells. The CsHT16 knockout mutant produces shorter fruits with a larger circumference, likely due to impaired sugar and phytohormone homeostasis. Concurrent reduction of CsHT3, CsHT12, and CsHT16 expression leads to decreased fruit size. Conversely, CsHT3 overexpression results in increased fruit size and higher fruit sugar levels. These findings suggest that CsHT16 plays an important role in maintaining sugar homeostasis, which shapes the fruit, while CsHT3, CsHT12, and CsHT16 collectively regulate the supply of carbohydrates required for cucumber fruit enlargement.