Received for publication 1 May 2019. Accepted for publication 17 June 2019. We thank Lesley Benyon, PhD, from Liwen Bianji, Edanz Group China (www.liwenbianji.cn/ac), for editing the English text of a draft of this manuscript. This work was supported by the National Technology System for Grape Industry (CARS-29-ZP-9) and National key research and development program (2018YFD0201300). 1These authors contributed equally. 2Corresponding author. E-mail: guoshunyang@ aliyun.com