国家自然科学基金项目31372507;青岛海洋科学与技术国家实验室鳌山科技创新计划项目2016ASKJ02;中央级公益性科研院所基本科研业务费项目20603022016001共同资助[This work was supported by the National Natural Science Foundation of China31372507;the Aoshan S&T Innovation Project from Qingdao National Laboratory of Marine Science and Technology2016ASKJ02;the Central Public-Interest Scientific Institution Basal Research Fund 20603022016001]