国家自然科学基金31461163005;中国水产科学研究院基本科研业务费2016HY-ZD02;山东省泰山学者攀登计划项目共同资助[This work was supported by the National Natural Science Foundation of China31461163005;Scientific Institution Basal Research Fund, Chinese Academy of Fishery Sciences2016HY-ZD02;Taishan Scholar Climbing Project, Shandong]