Hydrothermal systems at offshore Taiwan: Unique biological and geochemical characteristics
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摘要: 台湾地处西太平洋构造活动带，近海发育了多处热液活动，其中最典型的为龟山岛和绿岛热液体系。本文对海峡两岸在龟山岛和绿岛热液的地球化学特征以及周边生物体的响应的研究进展进行了综述。龟山岛热液喷出流体具有全球最低的pH(1.52)，富含重金属元素和CO2等酸性气体，是周围海水中有色溶解有机质的来源；绿岛具有全球热液中最低的溶解有机碳浓度(14 μmol /L)，且具有特殊的动力学特性。喷口周边分布了较为罕见的自然硫烟囱体和硫磺球。喷口的高毒性、高酸性热液改变了热液区生物体如螃蟹的生活习性和解毒机制。热液区的主要活跃菌群为参与碳、硫和氮代谢途径的γ-和ε-变形菌。主要生物质合成以硫还原和硫氧化的化能无机自养型生物为主，微生物硫代谢促进了热液系统中的微生物能量流动和元素循环作用。某些热液生物采用繁殖期迁徙的机制应对高毒性、高酸性热液环境。在热液活动的胁迫下，这些微生物产生了新颖独特的代谢产物。此外，龟山岛和绿岛热液体系还受到了潮汐、台风和地震等灾害性事件的影响。台湾近海热液体系的研究对认识热液地球化学循环、探讨热液的生态环境效应等具有重要的意义。Abstract: Offshore eastern Taiwan situated in the West Pacific active zone, there are multiple hydrothermal vents among which Kueishantao (KST) and Lutao (LT) host the most active systems. This paper reviewed the progress of the geochemical characteristics and biological responses of the KST and LT hydrothermal systems. The vent fluids emanated from the shallow KST hydrothermal vents have the world's lowest pH (1.52) values of any submarine vents. The fluids are rich in heavy metals and are accompanied by gases composed mostly by CO2. The hydrothermal vents are considered to be a source for chromophoric and fluorescent dissolved organic matter in the oceans. The hydrothermal systems showed the world’s lowest concentration of dissolved organic carbon (14 μmol/L) and unique kinetic characteristics in Lutao. Native sulfur chimneys and sulfur balls were found around the KST vents. The highly toxic and acidic KST vent fluids have disturbed the behavior of ambient macro-organisms including crabs which have developed detoxication measures. The active bacteria groups in the KST field are dominated by Gammaproteobacteria and Epsilonbacteraeota that are involved in the carbon, sulfur, and nitrogen metabolic pathways. Sulfur-reducing and sulfide-oxidizing chemolithoautotrophs account for most of the primary biomass synthesis, which fuels microbial energy flow and element cycling in the hydrothermal systems. Vent crab evolved an adaptive modulation of reproductive behavior to survive in the hydrothermal vent field. Under the stress of the KST hydrothermal activity, the microorganisms have generated cryptic compounds and metabolic products. Furthermore, both the KST and LD hydrothermal systems are affected by tides and catastrophic events such as typhoons and earthquakes. The studies on the hydrothermal systems at offshore Taiwan are of essential importance for investigating the geochemical cycles and eco-environmental impacts of global hydrothermal systems.1) Lebrato M, Garbe-Schönberg D, Tseng L C, et al. Earthquake and typhoon trigger shifts in shallow vents biogeochemistry analogous to human-made ocean disturbances[J]. Submitted to Scientific Reports, 2019.
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图 1 台湾近海构造地质背景（a）；龟山岛热液系统的远景和俯瞰图（b, c）；绿岛热液体系的典型喷口（d）
Fig. 1 Geological setting of offshore Taiwan (a); outlook of the Kueishantao hydrothermal system (b, c); and a typical vent of the Lutao geothermal field (d)
图 3 龟山岛热液独特的自然硫烟囱体(a)、硫磺丘(b)、硫磺球(c)和热液喷口周围生活的特有物种乌龟怪方蟹(d)
Fig. 3 the unique native sulfur chimney (a), sulfur mound (b), sulfur balls (c), and Xenograpsus testudinatus of the Kueishantao hydrothermal system (d)
图 4 龟山岛热液流体温度受到全日潮的影响，2000年台风“碧利斯”摧毁了所监测的喷口使其不再活动[4] (a)；潮汐对绿岛热液“户外池”水深的影响(b)
Fig. 4 The Kueishantao vent fluids were affected by diurnal tides, and the Typhoon “Bilis” at 2000 destroyed the monitored vent (a); semi-diurnal oscillation of water depth induced by tide of the Lutao hydrothermal system (b)
图 5 热液口微生物介导的生物地球化学过程，以及这些独特生态系统中碳、硫和氮循环的可能耦合机制（根据文献[23]重新绘制）
rTCA/CCB表示还原性三羧酸循环/卡尔文循环
Fig. 5 Microbial involved biogeochemical process, and the carbon, sulfur and nitrogen cycles and possible coupling mechanisms of this unique hydrothermal ecological system (modified from referance [23])
rTCA/CCB represents reductive tricarboxylic acid cycle/Calvin-Benson-Bassham cycle
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