国家自然科学基金项目41576147, 41306178;泰山领军人才工程高效生态农业创新类计划项目LJNY2015002;中国水产科学研究院基本科研业务费2018HY-ZD0201共同资助 [This work was supported by National Natural Science Foundation of China41576147, 41306178;Program of Shandong Leading TalentLJNY2015002;Central Public-Interest Scientific Institution Basal Research Fund, CAFS2018HY-ZD0201]