固氮蓝细菌的一种生物标志物——异形胞糖脂及其研究进展

doi:10.11867/j.issn.1001-8166.2019.09.0901

异形胞蓝细菌是重要的固氮微生物，广泛分布于淡水与海洋环境，其细胞壁内侧含有由糖苷基连接长链二醇、三醇、酮醇或酮二醇烷基而形成的糖脂，被称为异形胞糖脂（Heterocyst Glycolipids, HGs）。HGs是异形胞蓝细菌独特的生标化合物，其结构较为稳定，在淡水、海洋、微生物席及沉积物中均被检测到。对HGs生物地球化学研究起步较晚，但近年来进展较快，发现它们在异形胞蓝细菌的生物分类和环境示踪方面具有很大潜力。主要从生物及有机地球化学角度讲述蓝细菌异形细胞的分化机制和HGs结构性质，总结HGs在指示环境温度、异形胞蓝细菌群落演变、陆源有机质输入以及固氮活动中的应用情况，并对其在重建古海洋固氮活动以及古气候的重建等方面提出研究展望。

关键词: 异形胞蓝细菌, 异形胞糖脂, 固氮, 生物标志物

Heterocystous cyanobacteria are important diazotrophs commonly found in freshwater and ocean. They can produce special glycolipids known as Heterocyst Glycolipids (HGs) in their inner membrane layer. HGs are typically long-chain diols, triols, or hydroxyketones connected with a sugar moiety. HGs are unique biomarkers of heterocystous cyanobactria, which are relative stable and can be detected in lakes, marine systems, microbial mats and sediments. The study of HGs in biogeochemistry has not been carried out until recent years. HGs are proved to have great potential in indicating heterocystous cyanbacteria community and environment conditions. In this paper, the mechanism of cyanobacterial heterocyst differentiation and structure of HGs were introduced. In particular, emphasis was put on the application of HGs to infer environmental temperature, heterocystous cyanobacteria community, terrestrial organic input and nitrogen fixationin. The prospect of future study was also proposed.

Key words: Heterocystous cyanobacteria, Heterocyst glycolipids, Nitrogen fixation, Biomarker.

中图分类号:

P593

图1 基于细菌16S rRNA序列构建的蓝细菌系统发育进化树
罗马数字I-V为蓝细菌形态学分类（Subsection I~V）;黑色方框：异形胞固氮蓝细菌；N及灰色标识：非异形胞固氮蓝细菌；无标识：非固氮蓝细菌 ^{[

2
]}

Fig.1 The phylogenetic trees of cyanobacteria inferred from 16S rRNA
Roman numerals denote cyanobacteria subsections I~V. Black box: Heterocystous diazotropic cyanobacteria; Grey shadow and N: Nonheterocystous diazotrophic cyanobacteria; Non-shaded: Non-diazotrophic cyanobacteria ^{[

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]}

图2 异形胞蓝细菌Anabaena及相关机制示意图
(a) Anavaena藻丝 ^{[

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]}；(b) 异形细胞透射电摬图 ^{[

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]}；(c) 异形细胞分化示意图 ^{[

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]}；(d) 异形细胞与营养细胞物质交换示意图 ^{[

34
]}；GL, HGL：糖脂层；PS, HEP：多聚糖层；C：极性藻青藻颗粒；H：异形细胞

Fig.2 Anabaena and related metabolites model
(a) Filament of Anabaena ^{[

26
]};(b) Transmission electron micrograph of hetercyst ^{[

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]}; (c) Heterocyst formation diagram ^{[

27
]}; (d) Transport of metabolites between heterocyst and vegetative cell ^{[

34
]}. GL, HGL: Glycolipids layer; PS, HEP: Polysaccharide layer; C: Polar cyanophycin granule; H: Heterocyst

图3 HGs结构图^{[

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,

45
,

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]}

Fig.3 HGs structures^{[

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,

45
,

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]}

图4 西班牙淡水及极地环境样品中HG₂₆与温度呈现“S”型曲线关系^{[

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]}

Fig.4 HG₂₆ index vs. temperature in Spanish freshwaters and polar environment samples and fitted sigmoidal curve^{[

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]}

图5 波罗的海沉积柱中HGs分布，及其所反映的全新世以来的不同水文时期^{[

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]}

Fig.5 The distribution of HGs in a sediment core in Baltic Sea and related hydrographical phases over the Holocene^{[

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]}

图6 东地中海沉积柱中环境参数及HGs含量变化分布^{[

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]}
HGs含量增加与δ ¹⁵N的负偏同时发生，代表固氮活动的增强

Fig.6 The bulk properties and HGs contents of in a sediment core in the Eastern Mediterranean^{[

59
]}
The increased contents of HGs coincide with a decrease of δ ¹⁵N suggesting enhanced dinitrogen fixation

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A biomarker of Diazotrophic Cyanobacteria: Heterocyst Glycolipids and Their Research Progress