国家自然科学基金项目31172447,31572227;国家海洋公益性行业科研专项201405010;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;the National Nature Science Foundation of China31172447,31572227;the Marine Public Welfare Industry Research201405010;the Scientific and Technological Innovation Project Financially Supported by Qingdao National Laboratory for Marine Science and Technology2015ASKJ01