中国水产科学研究院基本科研业务费2016HY-ZD04;国家自然科学基金项目31372523;青岛海洋科学与技术国家实验室鳌山科技创新计划项目2015ASKJ01;泰山学者良种工程项目共同资助[This work was supported by Centeral Public-Interest Scientific Institution Basal Rasal Research Fund,CAFS2016HY-ZD04;National Natural Science Foundation of China31372523;the Scientific and Technological Innovation Project Financially Supported by Qingdao National Laboratory for Marine Science and Technology2015ASKJ01;Taishan Scholar Program for Seed Industry]