中国近海浮游植物光合溶解有机碳生产研究
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摘要
浮游植物光合作用合成的有机物，除以颗粒有机物的形态结合到细胞体中以外，还有相当一部分直接以溶解有机物(photosynthetically produced dissolvedorganic carbon, PDOC)的形态释放。然而以往绝大多数海洋生产力观测只对颗粒态部分的初级生产力进行测定，没有对PDOC进行同步测定。与此相应，大多数相关的食物网营养动力学模型，碳循环和碳收支模型也忽略了这一部分重要的碳流。由此造成对全球海洋初级生产力和碳吸收能力的低估。
本研究首次在中国黄海、长江口—东海、南海北部等典型海域开展系统的浮游植物光合溶解有机碳现场观测研究，利用改进的~(14)C同位素示踪方法获取海区的PDOC生产力分布资料以及光合有机碳细胞外释放比例(percentage ofextracellular release，PER)比的分布变化特征。研究除对中国近海浮游植物PDOC释放的环境调控机制，浮游植物物种特征与PDOC释放的关系，浮游植物PDOC释放对微食物环的作用等进行了系统分析外，并对浮游植物季节性水华、河口季节性低氧过程等特定生态过程与浮游植物PDOC释放之间的关系进行了讨论。主要结果如下。
中国各海区真光层初级生产力的PER比变化范围为7.6~39.3%，平均PER比为22.2%。据此结果对中国近海固碳量进行粗略估算，结果显示在把PDOC释放纳入计算的情况下中国近海总固碳量约为9.4×10~8t。忽略浮游植物溶解态光合产品而造成的中国近海固碳量低估约为2.5×10~8t。
对2009年黄海春季水华过程的现场观测研究显示，春季水华期浮游植物PDOC生产力与PER比均高于水华发生前。而在所观测的两个次表层水华过程中，以较小粒径的甲藻为优势物种的水华光合溶解有机碳释放明显高于以较大粒径硅藻为原因物种的水华过程。水华期浮游植物溶解有机物的大量释放，促进了海区异养浮游细菌生物量迅速上升，显示PDOC生产力对微食物环的重要意义。
对2006年6月-10月长江口及其毗邻海域浮游植物初级生产力的分布变化，及其与河口底层低氧的形成和发展之间的关系进行的研究结果显示，长江口夏季低氧区的形成与浮游植物生产力的变化密切相关，低氧核心区的形成与同一区域内大约20天前发生的河口强烈水华事件有直接关系。夏季高生产力期浮游植物的PDOC释放速率也出现高值，但真光层水体浮游植物溶解有机碳大量释放与底层低氧的出现并无直接关系。2006年8月低氧区的北移与2006年我国夏季出现大范围高温干旱天气，长江入海径流量大幅减少之间存在密切关系。
2009年夏季和冬季南海北部海区异养细菌分布及浮游植物PDOC生产力分布的对比分析结果显示，在寡营养外海区域，来自浮游植物直接释放的溶解有机物对细菌生产力的维持更为重要，而在近岸海区，更多来源的溶解有机物可供异养细菌利用，浮游植物释放的重要性相对降低。细菌需碳量估算结果显示，直接来自PDOC生产的溶解有机物不能满足南海北部异养浮游细菌的需要，浮游植物直接释放以外的溶解有机物供应对支持细菌生长也非常重要。
各海区PDOC环境调控机制综合分析结果显示，中国近海浮游植物光合溶解有机碳释放主要控制因子包括光照、营养盐水平和浮游植物粒径特征等。其中光照是控制PDOC生产垂直分布特征的重要因素，垂直分布上，PER高值经常出现在存在光抑制的表层和光照微弱的真光层底部水体。在水平分布上，外海以小粒径浮游植物占优势的寡营养海域PER显著高于大粒径浮游植物占优势的近海富营养海区。
Organic compounds synthesized by photosynthesis of phytoplankton, in additionto the part binding to the cell body in the form of the particulate organic matter, still aconsiderable part direct release in the form of photosynthetically produced dissolvedorganic carbon (PDOC). However, most past marine productivity observation wereonly determined on particulate portion of the primary productivity, no simultaneousdetermination on PDOC. Accordingly, most related food web trophic dynamics model,carbon cycle and carbon balance model also ignored this important part of carbonflow. This resulting in primary productivity and carbon sequestration capacityunderestimated in the global ocean.
This study first carried out systematic field observation on phytoplankton PDOCin typical areas such as the Yellow Sea, the Yangtze River Estuary-East China Sea,,northern South China Sea, using the improved14C isotope tracer method to obtain thePDOC productivity distribution data as well as the distribution and variationcharacteristics of PER (percentage of extracellular release) ratio in study areas. In thisstudy, in addtion to system analysis on environmental regulation mechanism of thephytoplankton PDOC release in China coastal sea, the relationship of phytoplanktonspecies characteristics and PDOC release, the effects of phytoplankton release onmicrobial food cycle, also discuss the relationship between phytoplankton PDOCrelease and specific ecological processes e.g. phytoplankton seasonal blooms, theseasonal hypoxia in estuaries. The main results are as follows.
PER ratio of primary productivity in the euphotic zone ranged from7.6to39.3%,the average PER was22.2%in different China sea. According to this result, carbonsequestration was roughly estimated in China coastal seas. The results showed thattotal carbon sequestration capacity was about9.4×10~8t in China coastal seas includingthe calculation of PDOC release. Ignoring phytoplankton PDOC products, carbonsequestration capacity in China coastal seas was underestimated about2.5×10~8t.
Field observation research to spring bloom processes in the Yellow Sea in2009showed that, phytoplankton PDOC productivity and PER ration in the spring bloom were higher than those before the occurrence of algal bloom. While in the observationof two subsurface water bloom processes, PDOC release was significantly higher inthe bloom process that dominant species was smaller size of dinoflagellates than thatwith reasons species as larger particle size diatom. Large amounts of phytoplanktonPDOC release in the bloom periods led to heterotrophic bacterioplankton biomassincreased rapidly in study areas, showing the significance of PDOC productivity tomicrobial food cycle.
The study results on the distribution variation of phytoplankton primaryproductivity in the Changjiang Estuary and its adjacent waters, and the relationshipwith the formation and development of hypoxia in the estuary bottom in Jun.-Oct.2006, showed that, the formation of the Yangtze River Estuary Hypoxia Zone insummer was closely related to the variation of phytoplankton productivity. Theformation of hypoxic core area had a direct relationship with strong algal bloomevents happened20days ago in the same area. In high productivity period in summer,phytoplankton PDOC release rate was high, but there was no direct relationshipbetween phytoplankton PDOC release in the euphotic layer and the underlyinghypoxia appear. There was close relationship between Hypoxia Zone movingnorthward in Aug.2006, and a wide range of high temperature and drought weather,Changjiang River Runoff substantially reduced in2006summer.
Research results on heterotrophic bacteria distribution and phytoplankton PDOCproductivity distribution in the northern South China Sea in2009summer and wintershowed that, relatively higher bacterial productivity appeared in offshore area thatwith a higher proportion of phytoplankton photosynthetic product release, illustratingphytoplankton PDOC release can more effectively used by heterotrophic bacterialproduction. But the results of bacterial carbon quantity estimation showed that, PDOCproduction can not meet the need of heterotrophic bacterioplankton in northern SouthChina Sea, supplied by other DOC outside of phytoplankton direct release is also veryimportant to bacterial growth.
Comprehensive analysis results on the environment control mechanism of PDOCshowed that, phytoplankton PDOC release controlled factors in China coastal seaincluding light, nutrients and phytoplankton size characteristics. The light was aimportant factor controlling the vertical distribution of PDOC production. In verticaldistribution, high PER ratio often occured in surface with the presence of lightinhibition and the euphotic layer bottom with week light. In horizontal distribution, PER in oligotrophic offshore waters dominated with small size phytoplankton weresignificantly higher than that in eutrophic coastal waters with large size phytoplanktondominant.
本研究首次在中国黄海、长江口—东海、南海北部等典型海域开展系统的浮游植物光合溶解有机碳现场观测研究，利用改进的~(14)C同位素示踪方法获取海区的PDOC生产力分布资料以及光合有机碳细胞外释放比例(percentage ofextracellular release，PER)比的分布变化特征。研究除对中国近海浮游植物PDOC释放的环境调控机制，浮游植物物种特征与PDOC释放的关系，浮游植物PDOC释放对微食物环的作用等进行了系统分析外，并对浮游植物季节性水华、河口季节性低氧过程等特定生态过程与浮游植物PDOC释放之间的关系进行了讨论。主要结果如下。
中国各海区真光层初级生产力的PER比变化范围为7.6~39.3%，平均PER比为22.2%。据此结果对中国近海固碳量进行粗略估算，结果显示在把PDOC释放纳入计算的情况下中国近海总固碳量约为9.4×10~8t。忽略浮游植物溶解态光合产品而造成的中国近海固碳量低估约为2.5×10~8t。
对2009年黄海春季水华过程的现场观测研究显示，春季水华期浮游植物PDOC生产力与PER比均高于水华发生前。而在所观测的两个次表层水华过程中，以较小粒径的甲藻为优势物种的水华光合溶解有机碳释放明显高于以较大粒径硅藻为原因物种的水华过程。水华期浮游植物溶解有机物的大量释放，促进了海区异养浮游细菌生物量迅速上升，显示PDOC生产力对微食物环的重要意义。
对2006年6月-10月长江口及其毗邻海域浮游植物初级生产力的分布变化，及其与河口底层低氧的形成和发展之间的关系进行的研究结果显示，长江口夏季低氧区的形成与浮游植物生产力的变化密切相关，低氧核心区的形成与同一区域内大约20天前发生的河口强烈水华事件有直接关系。夏季高生产力期浮游植物的PDOC释放速率也出现高值，但真光层水体浮游植物溶解有机碳大量释放与底层低氧的出现并无直接关系。2006年8月低氧区的北移与2006年我国夏季出现大范围高温干旱天气，长江入海径流量大幅减少之间存在密切关系。
2009年夏季和冬季南海北部海区异养细菌分布及浮游植物PDOC生产力分布的对比分析结果显示，在寡营养外海区域，来自浮游植物直接释放的溶解有机物对细菌生产力的维持更为重要，而在近岸海区，更多来源的溶解有机物可供异养细菌利用，浮游植物释放的重要性相对降低。细菌需碳量估算结果显示，直接来自PDOC生产的溶解有机物不能满足南海北部异养浮游细菌的需要，浮游植物直接释放以外的溶解有机物供应对支持细菌生长也非常重要。
各海区PDOC环境调控机制综合分析结果显示，中国近海浮游植物光合溶解有机碳释放主要控制因子包括光照、营养盐水平和浮游植物粒径特征等。其中光照是控制PDOC生产垂直分布特征的重要因素，垂直分布上，PER高值经常出现在存在光抑制的表层和光照微弱的真光层底部水体。在水平分布上，外海以小粒径浮游植物占优势的寡营养海域PER显著高于大粒径浮游植物占优势的近海富营养海区。
Organic compounds synthesized by photosynthesis of phytoplankton, in additionto the part binding to the cell body in the form of the particulate organic matter, still aconsiderable part direct release in the form of photosynthetically produced dissolvedorganic carbon (PDOC). However, most past marine productivity observation wereonly determined on particulate portion of the primary productivity, no simultaneousdetermination on PDOC. Accordingly, most related food web trophic dynamics model,carbon cycle and carbon balance model also ignored this important part of carbonflow. This resulting in primary productivity and carbon sequestration capacityunderestimated in the global ocean.
This study first carried out systematic field observation on phytoplankton PDOCin typical areas such as the Yellow Sea, the Yangtze River Estuary-East China Sea,,northern South China Sea, using the improved14C isotope tracer method to obtain thePDOC productivity distribution data as well as the distribution and variationcharacteristics of PER (percentage of extracellular release) ratio in study areas. In thisstudy, in addtion to system analysis on environmental regulation mechanism of thephytoplankton PDOC release in China coastal sea, the relationship of phytoplanktonspecies characteristics and PDOC release, the effects of phytoplankton release onmicrobial food cycle, also discuss the relationship between phytoplankton PDOCrelease and specific ecological processes e.g. phytoplankton seasonal blooms, theseasonal hypoxia in estuaries. The main results are as follows.
PER ratio of primary productivity in the euphotic zone ranged from7.6to39.3%,the average PER was22.2%in different China sea. According to this result, carbonsequestration was roughly estimated in China coastal seas. The results showed thattotal carbon sequestration capacity was about9.4×10~8t in China coastal seas includingthe calculation of PDOC release. Ignoring phytoplankton PDOC products, carbonsequestration capacity in China coastal seas was underestimated about2.5×10~8t.
Field observation research to spring bloom processes in the Yellow Sea in2009showed that, phytoplankton PDOC productivity and PER ration in the spring bloom were higher than those before the occurrence of algal bloom. While in the observationof two subsurface water bloom processes, PDOC release was significantly higher inthe bloom process that dominant species was smaller size of dinoflagellates than thatwith reasons species as larger particle size diatom. Large amounts of phytoplanktonPDOC release in the bloom periods led to heterotrophic bacterioplankton biomassincreased rapidly in study areas, showing the significance of PDOC productivity tomicrobial food cycle.
The study results on the distribution variation of phytoplankton primaryproductivity in the Changjiang Estuary and its adjacent waters, and the relationshipwith the formation and development of hypoxia in the estuary bottom in Jun.-Oct.2006, showed that, the formation of the Yangtze River Estuary Hypoxia Zone insummer was closely related to the variation of phytoplankton productivity. Theformation of hypoxic core area had a direct relationship with strong algal bloomevents happened20days ago in the same area. In high productivity period in summer,phytoplankton PDOC release rate was high, but there was no direct relationshipbetween phytoplankton PDOC release in the euphotic layer and the underlyinghypoxia appear. There was close relationship between Hypoxia Zone movingnorthward in Aug.2006, and a wide range of high temperature and drought weather,Changjiang River Runoff substantially reduced in2006summer.
Research results on heterotrophic bacteria distribution and phytoplankton PDOCproductivity distribution in the northern South China Sea in2009summer and wintershowed that, relatively higher bacterial productivity appeared in offshore area thatwith a higher proportion of phytoplankton photosynthetic product release, illustratingphytoplankton PDOC release can more effectively used by heterotrophic bacterialproduction. But the results of bacterial carbon quantity estimation showed that, PDOCproduction can not meet the need of heterotrophic bacterioplankton in northern SouthChina Sea, supplied by other DOC outside of phytoplankton direct release is also veryimportant to bacterial growth.
Comprehensive analysis results on the environment control mechanism of PDOCshowed that, phytoplankton PDOC release controlled factors in China coastal seaincluding light, nutrients and phytoplankton size characteristics. The light was aimportant factor controlling the vertical distribution of PDOC production. In verticaldistribution, high PER ratio often occured in surface with the presence of lightinhibition and the euphotic layer bottom with week light. In horizontal distribution, PER in oligotrophic offshore waters dominated with small size phytoplankton weresignificantly higher than that in eutrophic coastal waters with large size phytoplanktondominant.
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