国家自然科学基金青年基金项目31302179;泰山学者良种工程项目、山东省重点研发计划项目2016GSF115030;中国水产科学研究院黄海水产研究所中央级公益性科研院所基本科研业务费专项资金20603022016006共同资助[This work was supported by National Natural Science Foundation of China31302179;Taishan Scholar Program for Seed Industry,Key Research and Development Project of Shandong Province2016GSF115030;Central Public-Interest Scientific Institution Basal Research Fund,YSFRI,CAFS20603022016006]