生物标志物及其在生态系统研究中的应用

doi:10.17521/cjpe.2019.0139

植物生态学报 ›› 2020, Vol. 44 ›› Issue (4): 384-394.DOI: 10.17521/cjpe.2019.0139

所属专题：全球变化与生态系统；生态学研究的技术和方法

生物标志物及其在生态系统研究中的应用

冯晓娟^1,^2,^*(),王依云^1,²,刘婷¹,贾娟¹,戴国华¹,马田^1,²,刘宗广¹

¹中国科学院植物研究所植被与环境变化国家重点实验室, 北京 100093
²中国科学院大学资源与环境学院, 北京 100049

收稿日期:2019-06-06 接受日期:2019-08-22 出版日期:2020-04-20 发布日期:2019-10-21
通讯作者: 冯晓娟 ORCID:0000-0002-0443-0628
基金资助:
国家重点研发计划(2017YFC0503902);国家重点基础研究发展计划(973计划)(2015CB954201);中国科学院创新交叉团队项目(JCTD-2018-06)

Biomarkers and their applications in ecosystem research

FENG Xiao-Juan^1,^2,^*(),WANG Yi-Yun^1,²,LIU Ting¹,JIA Juan¹,DAI Guo-Hua¹,MA Tian^1,²,LIU Zong-Guang¹

¹State Key Laboratory of Vegetation and Environmental Change, Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China
²College of Resources and Environment, University of Chinese Academy of Sciences, Beijing 100049, China

Received:2019-06-06 Accepted:2019-08-22 Online:2020-04-20 Published:2019-10-21
Contact: FENG Xiao-Juan ORCID:0000-0002-0443-0628
Supported by:
National Key R&D Program of China(2017YFC0503902);State Key Development Program for Basic Research of China(2015CB954201);Chinese Academy of Sciences Interdisciplinary Project(JCTD-2018-06)

摘要/Abstract

摘要：

生物标志物是环境和地质体中记载着原始生物母质分子结构信息的有机化合物, 其含量可以指征特定生物来源对天然有机质的相对贡献, 其组成和同位素信息还可以记录有机质的转化及环境信息。与传统元素及组分分析相比, 生物标志物为研究天然有机质的来源、动态变化和转化特征提供了具有高度专一性和灵敏度的工具, 因此, 近年来被广泛地应用于生态学和生物地球化学研究中。特别是, 与生态系统观测以及控制实验相结合, 生物标志物在揭示微生物的活性与碳源变化、土壤有机碳的稳定机制及其对全球变化的响应等方面显示了广阔的应用前景。近些年开发的生物标志物单体同位素分析也在生态系统碳氮周转与食物网研究等方面显示了巨大的研究潜力。基于此, 该文综述了生态系统研究中常用的生物标志物的种类、分析方法和应用方向, 总结了生物标志物研究目前存在的问题, 并对未来的研究方向进行了展望, 旨在为使用生物标志物的生态学和环境科学研究者提供参考。

关键词: 生物标志物, 生态系统研究, 土壤有机质, 单体同位素分析, 碳循环, 食物网

Abstract:

Biomarkers are biogenic organic compounds that carry the chemical structures specific to their biological sources and survive long-term preservation in environmental and geological systems. The abundance of biomarkers may indicate the relative contribution of specific biological sources to the natural organic matter while their chemical and isotopic compositions may also inform on the transformation stage of organic matter and the environmental settings. Compared with conventional bulk analysis, biomarkers offer highly specific and sensitive tools to track the sources, transformation and dynamic changes of natural organic matter components and have therefore been widely used in ecological and biogeochemical studies in the past decades. In particular, combined with ecosystem observations and control experiments, biomarkers have shown great potentials in revealing changes in microbial activity and carbon sources, soil organic matter dynamics, stabilization mechanisms and response to global changes. The recently-developed biomarker-specific isotope analysis also exhibits a great promise in revealing ecosystem carbon and nitrogen turnover and food web structures. This review summarizes several major categories of commonly used biomarkers, their analytical methods, applications in ecosystem studies and existing pitfalls, and discusses future directions of research to provide guidance for biomarker users in ecology and environmental sciences.

Key words: biomarker, ecosystem research, soil organic matter, compound-specific isotope analysis, carbon cycling, food web

冯晓娟, 王依云, 刘婷, 贾娟, 戴国华, 马田, 刘宗广. 生物标志物及其在生态系统研究中的应用. 植物生态学报, 2020, 44(4): 384-394. DOI: 10.17521/cjpe.2019.0139

FENG Xiao-Juan, WANG Yi-Yun, LIU Ting, JIA Juan, DAI Guo-Hua, MA Tian, LIU Zong-Guang. Biomarkers and their applications in ecosystem research. Chinese Journal of Plant Ecology, 2020, 44(4): 384-394. DOI: 10.17521/cjpe.2019.0139

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2019.0139

https://www.plant-ecology.com/CN/Y2020/V44/I4/384

图/表 3

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生物标志物 Biomarker	分子组成 Molecular composition	提取方法 Extraction method	研究意义 Research implication	关键参数 Key parameter
植物蜡质脂类 Plant lipids	长链(>C₂₀)正构烷烃、脂肪酸、脂肪醇、固醇等 Long-chain (>C₂₀) n-alkanes, n-alkanoic acids, n-alkanols, steroids, etc.	溶剂萃取 Solvent extraction	代表植物来源的脂类 Indicating lipids from terrestrial vegetation	ACL, CPI
角质单体 Cutin monomers	短链(C₁₄-C₁₈)羟基-环氧酸 Short-chain (C₁₄-C₁₈) hydroxy- and epoxy acids	碱式水解 Base hydrolysis	代表叶片角质来源的脂类 Indicating lipids from leaf cuticles	ω-C₁₆/∑C₁₆, ω-C₁₈/∑C₁₈
软木脂单体 Suberin monomers	长链(C₂₀-C₃₂)脂肪酸、双酸、ω-羟基酸等 Long-chain (C₂₀-C₃₂) aliphatic acids, diacids and ω-hydroxy acids, etc.	碱式水解 Base hydrolysis	代表根系和树皮软木脂来源的脂类 Indicating lipids from suberin in roots and barks
磷脂脂肪酸 Phospholipid fatty acids (PLFAs)	短链脂肪酸(<C₂₀) Short-chain fatty acids (<C₂₀)	Bligh-Dyer	指征活体微生物生物量和群落结构 Indicating microbial biomass and community structure	F/B
甘油二烷基甘油四醚Glycerol dialkyl glycerol tetraethers (GDGTs)	类异戊二烯和支链GDGT Isoprenoid and branched GDGTs	溶剂萃取 Solvent extraction	古菌和细菌细胞膜脂 Indicating archaeal and bacterial membrane lipids	MBT, CBT
中性糖 Neutral sugars	葡萄糖、半乳糖、甘露糖、核糖、木糖和阿拉伯糖等 Glucose, galactose, mannose. ribose, xylose, arabinose, etc.	酸式水解 Acid hydrolysis	区分植物和微生物来源的糖类 Indicating plant- and microbe-derived sugars	GM/AX, RF/AX
木质素酚类 Lignin phenols	带甲氧基的酚类化合物 Phenolic compounds with methoxyl groups	碱性氧化铜氧化法 Alkaline CuO oxidation	指示维管束植物来源的有机质和木质素氧化程度 Indicating lignin inputs from vascular plants and lignin oxidation stage	S/V, C/V, (Ad/Al)_V, (Ad/Al)_S
氨基酸 Amino acids	天冬氨酸、谷氨酸、丙氨酸等 Aspartic acid, glutamic acid, alanine, etc.	酸式水解 Acid hydrolysis	微生物、动植物及其蛋白质的代谢产物 Indicating proteinaceous inputs from microbes, animals and plants	D/L
氨基糖 Amino sugars	氨基葡萄糖、氨基半乳糖、甘露糖胺、胞壁酸 Glucosamine, galactosamine, mannosamine, muramic acid	酸式水解 Acid hydrolysis	微生物细胞壁组分, 常指征环境中的微生物残体碳 Components of microbial cell wall; indicating microbial necromass in soils	GluN/MurN

生物标志物 Biomarker	分子组成 Molecular composition	提取方法 Extraction method	研究意义 Research implication	关键参数 Key parameter
植物蜡质脂类 Plant lipids	长链(>C₂₀)正构烷烃、脂肪酸、脂肪醇、固醇等 Long-chain (>C₂₀) n-alkanes, n-alkanoic acids, n-alkanols, steroids, etc.	溶剂萃取 Solvent extraction	代表植物来源的脂类 Indicating lipids from terrestrial vegetation	ACL, CPI
角质单体 Cutin monomers	短链(C₁₄-C₁₈)羟基-环氧酸 Short-chain (C₁₄-C₁₈) hydroxy- and epoxy acids	碱式水解 Base hydrolysis	代表叶片角质来源的脂类 Indicating lipids from leaf cuticles	ω-C₁₆/∑C₁₆, ω-C₁₈/∑C₁₈
软木脂单体 Suberin monomers	长链(C₂₀-C₃₂)脂肪酸、双酸、ω-羟基酸等 Long-chain (C₂₀-C₃₂) aliphatic acids, diacids and ω-hydroxy acids, etc.	碱式水解 Base hydrolysis	代表根系和树皮软木脂来源的脂类 Indicating lipids from suberin in roots and barks
磷脂脂肪酸 Phospholipid fatty acids (PLFAs)	短链脂肪酸(<C₂₀) Short-chain fatty acids (<C₂₀)	Bligh-Dyer	指征活体微生物生物量和群落结构 Indicating microbial biomass and community structure	F/B
甘油二烷基甘油四醚Glycerol dialkyl glycerol tetraethers (GDGTs)	类异戊二烯和支链GDGT Isoprenoid and branched GDGTs	溶剂萃取 Solvent extraction	古菌和细菌细胞膜脂 Indicating archaeal and bacterial membrane lipids	MBT, CBT
中性糖 Neutral sugars	葡萄糖、半乳糖、甘露糖、核糖、木糖和阿拉伯糖等 Glucose, galactose, mannose. ribose, xylose, arabinose, etc.	酸式水解 Acid hydrolysis	区分植物和微生物来源的糖类 Indicating plant- and microbe-derived sugars	GM/AX, RF/AX
木质素酚类 Lignin phenols	带甲氧基的酚类化合物 Phenolic compounds with methoxyl groups	碱性氧化铜氧化法 Alkaline CuO oxidation	指示维管束植物来源的有机质和木质素氧化程度 Indicating lignin inputs from vascular plants and lignin oxidation stage	S/V, C/V, (Ad/Al)_V, (Ad/Al)_S
氨基酸 Amino acids	天冬氨酸、谷氨酸、丙氨酸等 Aspartic acid, glutamic acid, alanine, etc.	酸式水解 Acid hydrolysis	微生物、动植物及其蛋白质的代谢产物 Indicating proteinaceous inputs from microbes, animals and plants	D/L
氨基糖 Amino sugars	氨基葡萄糖、氨基半乳糖、甘露糖胺、胞壁酸 Glucosamine, galactosamine, mannosamine, muramic acid	酸式水解 Acid hydrolysis	微生物细胞壁组分, 常指征环境中的微生物残体碳 Components of microbial cell wall; indicating microbial necromass in soils	GluN/MurN

生物标志物及其在生态系统研究中的应用

Biomarkers and their applications in ecosystem research

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