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引用本文:王广策,王辉,高山,郇丽,王旭雷,顾文辉,解修俊,张建恒,孙松,于仁成,何培民,郑阵兵,林阿朋,牛建峰,王立军,张宝玉,沈颂东,卢山.绿潮生物学机制研究.海洋与湖沼,2020,51(4):789-808.
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绿潮生物学机制研究
王广策,王辉,高山,郇丽,王旭雷,顾文辉,解修俊,张建恒,孙松,于仁成,何培民,郑阵兵,林阿朋,牛建峰,王立军,张宝玉,沈颂东,卢山
1.中国科学院海洋研究所 实验海洋生物学重点实验室 青岛 266071;2.青岛海洋科学与技术试点国家实验室 海洋生物学与生物技术功能实验室 青岛 266237;3.中国科学院海洋大科学研究中心 青岛 266071;4.上海海洋大学 海洋生态与环境学院 上海 201306;5.中国科学院海洋研究所 海洋生态与环境科学重点实验室 青岛 266071;6.苏州大学 医学部基础医学与生物科学学院 苏州 215123;7.南京大学 生命科学学院 南京 210023
摘要:
绿潮发生机制极其复杂,但其核心机理是绿潮暴发时漂浮“藻席”中的浒苔快速形成巨大生物量,涉及浒苔生物学过程对环境的响应及其生理生化基础。为此,本文在已有研究报道的基础上,对浒苔生活史研究进行了充分的理论补遗;分析了漂浮“藻席”中浒苔生长与繁殖过程,认为该过程是绿潮生物量形成的关键;阐述了浒苔孢子囊形成是对富营养化背景下海水溶解无机氮(DIN)中硝态氮高占比的响应,认为一氧化氮分子可促进浒苔营养细胞向孢子囊的转化;解析了浒苔细胞对逆境因子的响应途径与机制。基于已有的研究成果,本文对浒苔引发绿潮的生物学过程进行了理论推定。人为或自然因素使固着浒苔处于漂浮状态,形成小规模“藻席”;富营养化背景下海水DIN中硝态氮占比的升高赋予了浒苔巨大的繁殖潜能,漂浮过程溶解无机碳(DIC)“充裕”和“不足”两个阶段的交替以及食藻动物啃食产生的藻片段使孢子囊形成比例大幅提升;孢子原位萌发等使释放的孢子在“藻席”中获得了附着基,个体数目随之指数增长;结合漂浮浒苔的高生长速率,“藻席”规模不断扩大,短时间内形成巨大生物量。同时,本文还对今后的绿潮研究提出一些建议,认为啃食动物在浒苔生物量消长过程中发挥重要作用,在漂浮“藻席”系统中扮演着“生态引擎”的功能,同时系统阐明浒苔孢子的原位萌发等现象的生物学机理应该是未来绿潮的重要研究内容。
关键词:  绿潮  浒苔  漂浮浒苔“藻席”  生活史  DIN中硝态氮占比  环境胁迫
DOI:10.11693/hyhz20200300078
分类号:X55
基金项目:国家产业技术体系,CARS-50号;中国科学院海洋大科学研究中心重点部署项目,COMS2019Q02号;国家自然科学基金,41976097号,41676157号,41876163号。
STUDY ON THE BIOLOGICAL MECHANISM OF GREEN TIDE
WANG Guang-Ce1,2,3, WANG Hui1,2,3, GAO Shan1,2,3, HUAN Li1,2,3, WANG Xu-Lei1,2,3, GU Wen-Hui1,2,3, XIE Xiu-Jun1,2,3, ZHANG Jian-Heng4, SUN Song5,3, YU Ren-Cheng5,3, HE Pei-Min4, ZHENG Zhen-Bing1,2,3, LIN A-Peng1, NIU Jian-Feng1,2,3, WANG Li-Jun1,2,3, ZHANG Bao-Yu1,2,3, SHEN Song-Dong6,7, LU Shan8
1.Key Laboratory of Experimental Marine Biology, Institute of Oceanology, Chinese Academy of Sciences, Qingdao 266071, China;2.Laboratory for Marine Biology and Biotechnology, Pilot National Laboratory for Marine Science and Technology(Qingdao), Qingdao 266237, China;3.Center for Ocean Mega-Science, Chinese Academy of Sciences, Qingdao 266071, China;4.College of Marine ecology and Environment, Shanghai Ocean University, Shanghai 201306, China;5.Key Laboratory of Marine Ecology and Environmental Sciences, Institute of Oceanology, Chinese Academy of Sciences, Qingdao 266071, China;6.School of Biology &7.Basic Medical Sciences, Soochow University, Suzhou 215123, China;8.School of Life Sciences, Nanjing University, Nanjing 210023, China
Abstract:
The mechanism of green tide outbreak is very complex, and its core mechanism is the large biomass accumulation of Ulva prolifera in the floating "algal mat" in the outbreak of green tide, which involves the response of biological process of U. prolifera to the environment factors and its physiological and biochemical bases. Based on the existing research, we made a full theoretical supplement to the study on life history of U. prolifera. By analyzing the growth and reproduction process of U. prolifera in floating "algal mat", we consider that this process is the key to the sharp increase of green tide biomass. In addition, we believe that the formation of U. prolifera sporangium is a response to the high proportion of nitrate nitrogen in dissolved inorganic nitrogen (DIN) in seawater under the background of eutrophication, and nitric oxide can promote the transformation of U. prolifera vegetative cells to sporangia. In addition, the response pathway and mechanism of U. prolifera cells to stresses were analyzed. Based on the existing research results, we proposed a theoretical presumption of the biological process of green tide caused by U. prolifera. First, artificial or natural factors make the attached U. prolifera in a floating state, forming a small-scale "algal mat". Under the background of eutrophication, the proportion of nitrate nitrogen in DIN increased, which gave U. prolifera a great proliferation potential. Secondly, the ratio of sporangium formation was greatly increased by the alternation of "abundant" and "insufficient" phases of dissolved inorganic carbon (DIC) in the floating process and the algal fragments produced by herbivores. In situ germination of spores made the released spores obtain attachment base in the "algal mat", and the number of individuals increased exponentially. Finally, due to the high growth rate of the floating U. prolifera, the scale of "algal mat" kept expanding, forming huge biomass in a short time. Based on this understanding, we put forward some suggestions for the future study of green tide. We believe that the gnawing animals play an important role in the growth and decline of U. prolifera biomass as an "ecological engine" in the floating "algal mat" system. At the same time, we think that systematic study on the biological mechanism of in situ spore germination of U. prolifera should be an important direction of future green tide research.
Key words:  green tide  Ulva prolifera  floating algal mat  life cycle  proportion of nitrate nitrogen in DIN  environmental stresses
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