从土壤腐殖质分组到分子有机质组学认识土壤有机质本质

doi:10.11867/j.issn.1001-8166.2019.05.0451

地球科学进展 ›› 2019, Vol. 34 ›› Issue (5): 451 -470. doi: 10.11867/j.issn.1001-8166.2019.05.0451

综述与评述下一篇

从土壤腐殖质分组到分子有机质组学认识土壤有机质本质

潘根兴(

),丁元君,陈硕桐,孙景玲,冯潇,张晨,郑聚锋,张旭辉,程琨,刘晓雨,卞荣军,李恋卿

南京农业大学资源环境科学学院土壤学系/农业资源与生态环境研究所，江苏南京 210095

收稿日期:2018-12-18 修回日期:2019-04-10 出版日期:2019-05-10
基金资助:
国家自然科学基金项目“秸秆生物质炭对农田土壤有机碳保持作用及其机制”（编号：41371298）和“稻田土壤有机碳固定与稳定化过程与机制”(编号：40830528)

Exploring the Nature of Soil Organic Matter from Humic Substances Isolation to SOMics of Molecular Assemblage

Genxing Pan( ),Yuanjun Ding,Shuotong Chen,Jingling Sun,Xiao Feng,Chen Zhang,Doross Marios,Jufeng Zheng,Xuhui Zhang,Kun Cheng,Xiaoyu Liu,Rongjun Bian,Lianqing Li

Department of Soil Science, and Institute of Resources, Ecosystem and Environment of Agriculture, Nanjing Agricultural University, Nanjing 210095, China

Received:2018-12-18 Revised:2019-04-10 Online:2019-05-10 Published:2019-07-04
About author:Pan Genxing(1958-), male, Nanjing City, Jiangsu Province, Professor. Research areas include soil organic matter and carbon sequestration. E-mail: gxpan@njau.edu.cn
Supported by:
Project supported by the National Natural Science Foundation of China “Effect of straw biochar on organic carbon preservation of croplands and the mechanism (No.43171298) and “Processes and mechanisms of C sequestration and stabilization in rice paddies of China: A study of soil-crop-microbes interactions”(No. 40830528)

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梳理了与土壤生态系统功能相联系的，特别是对固碳减排的土壤有机质本质认识的研究进展及路径，探讨了经典腐殖质学说存在的问题，概述了新近的有机质保护稳定学说及腐殖质组学学说，并追溯了生物标志物有机质分子研究，最后从土壤学的基本理念和理论出发讨论和重新认识土壤有机质的本质及其价值。从形成条件、分离条件和分子鉴定等多方面分析，土壤腐殖质形成和稳定学说越来越显示出局限性；而面向气候变化的碳固定研究可以深入探析土壤有机质的复杂存在状态。越来越认识到土壤有机质是投入土壤的有机物质经不同程度生物利用或降解的产物残留，只是被土壤不同程度地区隔和封闭，本质上仍是分子量变化极大的生命源有机物的集合。因此，可通过生物标志物分子作为靶标在土壤中提取和识别，该技术的发展将孕育萌生土壤有机质分子组学。后者可以用于判读土壤有机质的结构支撑、反应活性和促生功能等方面的本质差别，这些差别可能是由有机分子组成结构及存在状态所决定而不是由有机分子稳定性决定的。从这个概念出发，类似于土壤微生物分子生态，土壤有机质的丰度、组成、结构与功能间的联系可能是土壤有机质本质的核心问题。对这种关系的量化和参数化表征可用以探索土壤有机质永续固定，且可以保持生命活性的土壤有机质的管理策略及技术，并配合土壤的团聚体理论诠释土壤的本质和生态系统功能服务，这将是未来土壤学服务人类可持续发展的理论立足点。

关键词: 土壤有机质, 土壤腐殖质, 腐殖质组学, 生物标志物, 分子有机质组学, 结构功能关系

In this review, the evolution of Soil Organic Matter (SOM) research was traced back to outline the main achievement of understanding SOM in relation to its ecological functioning, particularly of carbon sequestration against climate change. The short-coming of soil humus theory, knowledge of SOM protection and stabilization, framework of newly emerged Humeomics as well as the increasingly active study of molecular organics in soils were analyzed and discussed, highlighting the importance of re-visiting SOM in term of structure-property-functions for the main mission of modern soil science. There were limitations of soil forming conditions, fraction separation procedure and single molecule identification for understanding the huge complex humus of larger sized synthesized molecules. Thanks to the ever-active studies of soil (organic) carbon sequestration and stabilization focusing on the association status of SOM with soil components, SOM has been increasingly recognized as an assemblage of metabolites from life activities on or in soil, with different allocation or protected in mineral/organic complex phases, which could be traced by biomarker molecules. Using such biomarker molecules as a target (like primer in molecular microbiology), all the molecules of SOM could be digested and isolated for qualitative or quantitative identification with GC/MS high resolution technologies. Such development has emerged a new paradigm of molecular SOM study, finally as SOMics as a modern soil science frontier. The functioning of SOM for stabilizing soil structure, enhancing reactivity and promoting biological resistance could be correlated to the paradigm of abundance, composition, structure and functions rather than the content and recalcitrance of SOM. This may deserve urgent studies to quantify and parameterize the defined paradigm based on the molecular composition of SOM. Again, such theory and technology development could provide a tool to manage SOM in term of carbon sequestration but revalorizing bioactivity in ecosystems, especially in agroecosystems. We believe such studies could rather depict the nature of SOM and of soil in relation to its ecological services and functioning, which will be the focus of soil science in serving the sustainable development of human society.

Key words: Soil organic matter, Humus, Humeomics, Biomarker molecules, SOMics, Structure-function relationship.

中图分类号:

P934

图1 土壤有机质概念及组成认识(a)^{[

62
]}和（b）^{[

63
]}的演变

Fig.1 Concept update of soil organic matter and the composition (a）^{[

62
]}和（b）^{[

63
]}

图2 土壤腐殖质组学的连续提取分离法程序框架（据参考文献[ 81 ]修改）

Fig.2 Procedure framework of sequential extraction and separation of soil humus（modified after reference [ 81 ]）

图3 腐殖质组学分离鉴定法获得的某水稻土土壤有机质组分Van Krevelen图

Fig.3 Van Krevelen plot of soil organic matter composition in a paddy soil obtained by Humeomics identification

表1 土壤有机质中可鉴定的生物标志物特征与源解析

Table 1 Characterization and source analysis of identifiable biomarkers

化合物类型	分子类别	碳链范围和官能团	生命体来源
Triterpenoids		β-amyrin, oleanolic acid, ursolic acid	被子植物生物质
Aliphatic lipids	n-alkanes	C₂₅-C₃₃ odd	高等植物蜡质、真菌生物质
	n-alkanoic acids	C₁₂-C₃₂ even	木栓质、角质、真菌生物质
	n-alkanoic acids	C₁₆-C₃₂ even	高等植物蜡质
	n-alkanols	C₁₆-C₃₂ even	高等植物蜡质、木栓质
	α, ω-alkanedioic acids	C₁₆, C₁₈	角质
	α, ω-alkanedioic acids	C₁₆-C₂₈ even	木栓质或木栓质结合的蜡质
	iso-alkanoic acids	C₁₆, C₁₈	细菌和真菌生物质
	ω-hydroxyalkanoic acids	C₁₆, C₁₈	木栓质、角质
	ω-hydroxyalkanoic acids	C₂₀-C₃₀ even	木栓质、高等植物蜡质
	Acylglycerides	C₁₆, C₁₈ monoacylglycerides	各类生物质的生物质
	di-, trihydroxyalkanoic acids	C₁₆, C₁₈	角质、木栓质
	α-hydroxyalkanoic acids	C₁₆, C₂₂-C₂₆ even	植物或者真菌生物质
Carbohydrates		glucose, mannose, sucrose	各类生物质
Carbohydrates		trehalose	真菌，也可能是植物生物质
Phenols		vanillin, acetovanillone, vanillic acid	木质素
		syringaldehyde, acetosyringone, syringic acid
		ferulic acid, p-coumaric acid
		ferulic acid, p-coumaric acid, vanillin	木栓质或木栓质结合的蜡质
		Syringaldehyde	木质素和纤维素混合体物
		ferulic acid,p-coumaric acid	木质素和纤维素混合体物
Steroids		β-sitosterol, stigmasterol,campesterol	木质素
		ergosterol	真菌
		cholesterol	动物、植物和真菌

表1 土壤有机质中可鉴定的生物标志物特征与源解析

Table 1 Characterization and source analysis of identifiable biomarkers

化合物类型	分子类别	碳链范围和官能团	生命体来源
Triterpenoids		β-amyrin, oleanolic acid, ursolic acid	被子植物生物质
Aliphatic lipids	n-alkanes	C₂₅-C₃₃ odd	高等植物蜡质、真菌生物质
	n-alkanoic acids	C₁₂-C₃₂ even	木栓质、角质、真菌生物质
	n-alkanoic acids	C₁₆-C₃₂ even	高等植物蜡质
	n-alkanols	C₁₆-C₃₂ even	高等植物蜡质、木栓质
	α, ω-alkanedioic acids	C₁₆, C₁₈	角质
	α, ω-alkanedioic acids	C₁₆-C₂₈ even	木栓质或木栓质结合的蜡质
	iso-alkanoic acids	C₁₆, C₁₈	细菌和真菌生物质
	ω-hydroxyalkanoic acids	C₁₆, C₁₈	木栓质、角质
	ω-hydroxyalkanoic acids	C₂₀-C₃₀ even	木栓质、高等植物蜡质
	Acylglycerides	C₁₆, C₁₈ monoacylglycerides	各类生物质的生物质
	di-, trihydroxyalkanoic acids	C₁₆, C₁₈	角质、木栓质
	α-hydroxyalkanoic acids	C₁₆, C₂₂-C₂₆ even	植物或者真菌生物质
Carbohydrates		glucose, mannose, sucrose	各类生物质
Carbohydrates		trehalose	真菌，也可能是植物生物质
Phenols		vanillin, acetovanillone, vanillic acid	木质素
		syringaldehyde, acetosyringone, syringic acid
		ferulic acid, p-coumaric acid
		ferulic acid, p-coumaric acid, vanillin	木栓质或木栓质结合的蜡质
		Syringaldehyde	木质素和纤维素混合体物
		ferulic acid,p-coumaric acid	木质素和纤维素混合体物
Steroids		β-sitosterol, stigmasterol,campesterol	木质素
		ergosterol	真菌
		cholesterol	动物、植物和真菌

图4 土壤有机质组分及其结构—组成—功能联系的连续分布概念框架

Fig.4 Conceptual framework of continuous distribution of soil organic matter components and their structure-constitution-function relationships

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