Biogeochemistry characteristic and contribution to carbon transportation of Transparent Exopolymer Particles(TEP) in the ocean
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摘要: 透明胞外聚合颗粒物（Transparent Exopolymer Particles，TEP）是在海洋中广泛存在的一类特殊的胞外聚合物，具有透明、含碳量高、黏性强等特点，兼具胶体和颗粒物的特性，在海洋碳输运过程中起重要作用。本文系统梳理了海洋中TEP的生物地球化学特征及其对海洋碳输运的影响。海洋中的TEP呈不定型状态，尺寸差异极大，主要由浮游植物所释放的前体物质自组装生成，并通过生物代谢、海气交换及沉降等过程从海洋中移除，其丰度和分布受浮游植物和细菌等生物的活动、海水理化环境以及动力过程的共同驱动。近岸海域中TEP的浓度变化范围极广（0-
14800 μg XG eq /L），通常随离岸距离的增加而降低且与生产力密切相关，因此呈现显著的季节变化；开阔大洋中TEP的浓度一般在0-200 μg XG eq /L，在上层水体中通常随深度的增加而降低，而在中深层水体分布较均匀。作为颗粒有机碳的重要组分，TEP占颗粒有机碳库的比例及其对颗粒有机碳沉降通量的贡献，在近海通常在40%以下，而在开阔大洋则高达50%以上。除了改变颗粒沉降速率影响碳垂直输送，TEP还能通过微表层的富集效应介导海气碳交换，但该过程的机理及对碳输运的贡献仍需进一步明确。未来研究应聚焦TEP定量检测方法、组成形态解析及其碳转换因子的优化，在此基础之上进一步阐释海洋中TEP的生物地球化学行为及其影响海洋碳输运的作用机制。Abstract: Transparent exopolymer particles (TEP) are a special class of extracellular polymeric substances which is ubiquitous in seawater, characterized by transparency; high carbon content and stickiness; with colloidal and particulate characteristics, play a significant role in marine carbon transportation. In this review, we describe the Biogeochemistry characteristic and contribution to carbon transportation of Transparent Exopolymer Particles (TEP) in the ocean systematically. Marine TEP are amorphous, their size is variable, mainly generated by self-assembly of precursors which is released by phytoplankton and removed from ocean through processes such as biological metabolism, air-sea exchange and settlement, their abundance and distribution driven by the activity of organisms such as phytoplankton such as phytoplankton and bacteria, as well as the physicochemical environment and hydrodynamic processes. TEP concentrations in coastal waters exhibit considerable variability(0-14,800 μg XG eq L-1) and decrease with increasing offshore distance and closely related to productivity, thus demonstrating marked seasonal variations; In the open ocean, TEP concentrations are usually lower (0-200 μg XG eq L-1) and decrease with depth in epipelagic waters, while remaining relatively steady in Mesopelagic and Bathypelagic waters. As an important component of particulate organic carbon (POC), TEP typically account for less than 40% of the POC pool and its sinking flux in coastal areas, but can contribute more than 50% in the open ocean. In addition to altering particle sinking rates and affecting vertical carbon transport, TEP can also mediate air-sea carbon exchange through enrichment in the sea surface microlayer, though the underlying mechanisms and their significance in carbon transport require further investigation. Future research should focus on improving quantitative detection methods for TEP, analyzing their composition and morphology, and optimizing carbon conversion factors. to better understand the biogeochemical behaviors of TEP and their mechanisms in marine carbon transportation. -
图 1 海洋中TEP的生物地球化学循环及驱动因素
Fig. 1 Biogeochemical cycle and driving factors of marine TEP
表 1 TEP的不同测定方法比较
Tab. 1 Comparison of different methods for TEP determination
方法 染色 检测限（μg XG eq/L） 使用硫酸 盐分干扰 分析时间 特点 参考文献 显微镜计数法 是 − 否 否 慢 可观察尺寸形态但费时费力 [1] 传统比色法 是 5 是 否 >2 h 操作简便但复现性低 [6] 快速分光光度法 是 100 否 是 ～30 min 速度较快但不适用海水 [7] 比色法 是 − 否 否 海水测量时间较长 可测量TEP前体，但海水需预处理 [8] TOC校准法 是 − 是 否 >2 h 结果精确但易受有机碳影响且单位不同 [11] 10kDa法 是 0.05 是 否 >2 h 可测量前体，复现性高且检测限低但步骤繁琐 [9] AB-DI法 是 5 否 否 >1 h 避免使用硫酸，TEP及其前体使用同一标准曲线 [10] UV254/光谱法 否 0.6-32或0.01-0.1 否 否 快速或>1 h 可快速测定但需预制标准曲线 [10] 顺磁性功能化微球体法 否 − 否 否 >160 min 无需染色和硫酸提取但存在损失且单位不同 [13] 在线监测法 是 5000 否 否 30 min-1 h 分析时间较短但检测限非常高 [14] 
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表 2 不同海域中Chl a及TEP的丰度
Tab. 2 Abundance of Chl a and TEP across Different oceans
海域 季节 采样深度（m） Chl a(mg/m3) TEP含量（μg XG eq/L） 文献 河口和海湾 切萨皮克湾河口 全年 0-23.7 − 492（37- 2820 ）[22] 圣劳伦斯河口 夏秋 0-50 − 15- 1548 （291）[6] 吉雄河口 夏 0-4 0.18-25.00 225- 1059 [23] 珠江口 夏冬 0-30 2.01-124.11 908.1（88.7- 1586.9 ）[24] 长江口 春夏秋 0-80 0.19-40.20 37.9- 1226.11 [25] 胶州湾 全年 0.5m 1.28±1.48 75-553 [26] 贾兰湾 全年 0-20 1.8±1.6 215.9±172.2（26.5- 1695.4 ）[27] 布拉内斯湾 全年 0.5-20 0.48±0.23 81.7±11.7（11.3-289.1） [28] 东京湾 全年 0-20 <5-81.2 14- 1774 [29] 加的斯湾 夏 5-100 − 0-600 [30] 近海 阿尔沃兰海 夏 0-75 − <25-539 [31] 波罗的海 夏 4-20 − 145-322 [32] 白令海 夏 0- 3421 − 34-628 [33] 爱琴海 春夏秋 0-100 0.19±0.12 73.1±35.2（15.4-188） [34] 亚得里亚海 全年 0-35 − 570（4- 14800 ）[35] 罗斯海 冬 0-150 4.0 308（0- 2800 ）[36] 地中海 春 3- 3164 <0.92 0.6-81.7 [37] 南海 夏 5- 3700 0-1.96 0.6-78.6 [38] 东海 全年 3-150 <1-17.14 115±67（28-376） [39] 闽东沿海 夏 0-40 0.61-18.95 201.8±177.9（25.2-935.5） [40] 温州近海 夏秋 0-150 1.47±2.11 0- 1374.7 [41] 开阔大洋 东北大西洋 春 0-70 0.07-0.59 10-124 [42] 太平洋 冬 0-200 0.09-1.4 25±7.4（5.3-40） [43] 西北太平洋 春夏 5-300 0.05-0.27 41.8±9.1（18-69） [44] 南大洋 夏 0-200 0.01-5.36 15.4±10（0-48.9） [45] 西北冰洋 秋 0-200 0.12-1.2 120±22（37-130） [43] 东南印度洋 春 0-200 0.18±0.16 20±6（5-46.4） [46]
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表 3 不同海域中TEP对POC贡献的比较
Tab. 3 Comparison of TEP Contribution to POC across different oceans
海域 季节 采样深度 POC(mg/L) 转换因子 POC% 文献 河口和海湾 切萨皮克湾河口 全年 0-23.7 − 0.53 32±16%（0-89%） [22] 纽斯河口 全年 表 − 0.75 16±9% [21] 贾兰湾 全年 0-20 0.40±0.19 0.51 2.4-78% [27] 多纳保拉湾 全年 1 − 0.75 6.9±5.8%（0.1-22.8%） [75] 胶州湾 全年 0.5 0.60±0.30 0.75 26% [26] 特拉诺瓦湾 春 表 0.09 0.51 9.8±3.1% [51] 夏 0.17 21.9±7.1% 秋 0.08 26.9±6.1% 冬 0.07 13.0±4.2% 近海 爱琴海东北岸 春 0-100 0.06±0.02 0.51 73.6±30.9%（37.5-147%） [34] 夏 0.05±0.02 79.7±27.7（33.3-136%） 秋 0.03±0.01 52.0±24.3%（14.4-111%） 东海沿海 夏 表-底 0.08-1.47 0.53 32.03% [76] 闽东沿海 夏 0-40（表-底） 0.24±0.09 0.53 44.8±101.7%（9.8-110.8%） [40] 温州近海 夏 0-150（表-底） 0.17±0.18 0.53 42.2% [41] 秋 0.65±1.09 41.90% 开阔大洋 东北大西洋 春 表 0.04±0.01 0.75 85.2±1.6%（27.2-139.2%） [37] 中 0.01 78.7±3.0%（33.5-182.8%） 深 0.01 66.7±1.2%（22.1-169.1%） 大西洋 秋 4 16.6±15.8 0.51 73±36%（28-110%） [65] 西北冰洋 秋 0-200 0.04 0.51 160% [43] 太平洋 冬 0-200 0.02 80%
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