地球科学进展

地球科学进展 ›› 2026, Vol. 41 ›› Issue (1): 25 -39. doi: 10.11867/j.issn.1001-8166.2026.008

综述与评述 上一篇    下一篇

美拉德反应：土壤有机碳库稳定过程中被忽视的非生物驱动机制
薛华敏1,2(), 张仲胜1(), 赵雯雯1,2, 于子成1, 武海涛1, 姜明1   
  1. 1.中国科学院东北地理与农业生态研究所，吉林 长春 130102
    2.中国科学院大学，北京 100049
  • 收稿日期:2025-10-20 修回日期:2025-12-25 出版日期:2026-01-10
  • 通讯作者: 张仲胜 E-mail:xuehuamin@iga.ac.cn;zzslycn@iga.ac.cn
  • 基金资助:
    国家自然科学基金重大项目(42494825)

Maillard Reaction: An Overlooked Abiotic Driver in the Formation and Stabilization of Soil Organic Carbon Pools

Huamin Xue1,2(), Zhongsheng Zhang1(), Wenwen Zhao1,2, Zicheng Yu1, Haitao Wu1, Ming Jiang1   

  1. 1.Northeast Institute of Geography and Agroecology, Chinese Academy of Sciences, Changchun 130102, China
    2.University of Chinese Academy of Sciences, Beijing 100049, China
  • Received:2025-10-20 Revised:2025-12-25 Online:2026-01-10 Published:2026-03-10
  • Contact: Zhongsheng Zhang E-mail:xuehuamin@iga.ac.cn;zzslycn@iga.ac.cn
  • About author:Xue Huamin, research areas include carbon and nitrogen cycling and biogeochemical processes in wetland ecosystems.E-mail: xuehuamin@iga.ac.cn
  • Supported by:
    the National Natural Science Foundation of China(42494825)
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在全球变化背景下，由岩石风化—矿物催化—非酶促聚合联动的美拉德反应，可能是连接矿物与土壤有机碳封存的地球化学桥梁，成为实现碳中和重要的潜在地球化学途径。美拉德反应在土壤中被证实可诱导还原糖和氨基化合物在路易斯酸的催化下发生缩合—聚合反应，生成结构稳定、难降解的类黑精等聚合物，成为重要的碳固存补充机制。美拉德反应依赖于高温、高pH和适度水分条件，其反应速率并显著受铁/锰氧化物和层状硅铝酸盐等矿物催化增强，进而在深层土壤和厌氧沉积物中有效提升土壤有机碳稳定性。美拉德反应产物（以类黑精结构为代表的一大类化合物）不仅自身具有生物惰性，更能通过与矿物形成物理限域、官能团络合及金属桥联的有机—无机复合体，显著提升碳持久封存能力。同时，美拉德反应作为连接植物来源有机碳、微生物转化产物与矿物稳定化过程的关键非酶促转化环节，丰富和发展了土壤腐殖质的形成机制。然而，天然土壤环境中美拉德反应研究尚处于起步阶段，研究多依赖模拟实验，缺乏原位观测数据与机制模型，尚未解析土壤中美拉德反应的特异性分子标志物，无法量化其在稳定性土壤有机碳形成中的贡献，多因子调控美拉德反应的交互效应与协同机制尚不清晰。提出美拉德反应通过“矿物界面催化—类黑精生成—有机矿物复合体形成”三步稳定化路径，为土壤惰性碳库的形成提供了超越传统微生物主导理论的新机制，填补了该领域对非生物化学过程认知的空白。未来应重点解析美拉德反应在“气候变化—岩石风化—碳稳定性”链条中的枢纽作用，将美拉德反应介导的非生物固碳模块纳入陆地碳汇模型中，为地质碳封存与生态碳汇协同增效技术研发提供机理支持。

关键词: 美拉德反应, 非酶促聚合, 类黑精, 路易斯酸, 矿物, 碳封存

The sequestration of Soil Organic Carbon (SOC) constitutes a pivotal component of global climate change mitigation strategies. While the “microbial carbon pump” and biotic anabolism have traditionally dominated conceptual paradigms of humification, emerging evidence suggests that this biocentric view may underestimate the contribution of abiotic geochemical pathways. This review systematically delineates the role of the non-enzymatic Maillard reaction as a critical geochemical bridge linking mineral weathering processes to long-term SOC persistence.We propose a mechanistic framework of “mineral interfacial catalysis-melanoidin formation-organo-mineral complexation”to elucidate this abiotic stabilization trajectory. Specifically, soil minerals, particularly Fe/Mn oxides and phyllosilicates, act as natural catalysts by providing Lewis acid sites that lower the apparent activation energy for the condensation and polymerization of reducing sugars and amino compounds. This process transforms labile precursors into chemically recalcitrant, aromatic-rich polymers commonly referred to as melanoidin-like substances. These reactions are thermodynamically and kinetically favored under conditions of elevated temperature, alkaline to neutral pH, and intermediate to fluctuating moisture regimes, potentially representing a dominant stabilization pathway in subsurface horizons or anaerobic environments where microbial activity is energetically or kinetically constrained.The resulting melanoidin-type products exhibit a pronounced dual-protection capacity, beyond their inherent structural heterogeneity and low biological accessibility, they form robust associations with mineral matrices through physical confinement, ligand exchange, and polyvalent cation bridging. By acting as a non-enzymatic hub that integrates plant-derived carbon inputs and microbial metabolites into persistent organo-mineral complexes, the Maillard reaction challenges conventional theories of humic substance formation and provides a mechanistic framework for an underexplored abiotic pathway of carbon stabilization.Despite growing recognition of its potential importance, current understanding remains constrained by a reliance on simplified laboratory proxies, a scarcity of in situ field evidence, and the absence of diagnostic molecular biomarkers capable of distinguishing abiotic melanoidins from microbially derived necromass. Consequently, the quantitative contribution of this abiotic module to long-term SOC persistence remains poorly constrained. Future research should prioritize resolving environmental controls, multi-factor interactions, and identifying the molecular fingerprints of soil Maillard products in natural ecosystems. Incorporating mineral-mediated, non-enzymatic stabilization processes into terrestrial carbon cycle models will be essential for accurately capturing the coupled “climate change-rock weathering-carbon stability”continuum and for informing strategies that integrate geological and ecological carbon sequestration.

Key words: Maillard reaction, Non-enzymatic polymerization, Melanoidin, Lewis acid, Minerals, Carbon sequestration

中图分类号: 

图1 美拉德反应机理［6］
Fig. 1 Mechanistic illustration of the Maillard reaction［6］
图2 土壤中美拉德反应发生示意图［4］
土壤中美拉德反应分为3个阶段，第一阶段为还原糖与胺类化合物之间的亲核缩合，由于矿物提供的路易斯酸点位可配位活化羰基碳，增强其亲电性，进而显著促进还原糖与氨基化合物之间的亲核加成产生席夫碱（Schiff base）及其Amadori重排产物；第二阶段主要包括糖裂解、羰基化合物生成及进一步的重排与氧化过程；第三阶段为产物间缩合、芳构化及聚合反应，最终形成分子量大、深色、结构复杂、高度芳香化且富含C-N/C=O功能团的类黑精。
Fig. 2 Schematic representation of the Maillard reaction occurring in soil environments［4］
The Maillard reaction in soil proceeds in three stages. The first stage involves nucleophilic condensation between reducing sugars and amine compounds. The Lewis acid sites provided by minerals can coordinate and activate the carbonyl carbon， enhancing its electrophilicity and significantly promoting the nucleophilic addition between reducing sugars and amino compounds to produce Schiff bases and their Amadori rearrangement products. The second stage mainly includes sugar cleavage， carbonyl compound formation， and further rearrangement and oxidation processes. The third stage involves condensation， aromatization， and polymerization reactions between the products， ultimately forming melanoidins—large， dark-colored molecules with complex structures， high aromaticity， and rich in C-N/C=O functional groups.
图3 水分活度和pH对美拉德反应速率的影响
（a）水分活度对美拉德反应速率的影响［40］；（b）pH对甘氨酸氨基有效浓度的影响（对美拉德反应速率的影响）［11］。
Fig. 3 Effects of water activity and pH on the rate of the Maillard reaction rate
（a） Influence of water activity on the Maillard reaction rate［40］； （b） Effect of pH on the effective concentration of glycine amino groups， impacting the Maillard reaction rate［11］.
图4 Birnessite催化作用下葡萄糖和甘氨酸美拉德反应的吸光度与波长关系图［20］
Fig. 4 Absorbance spectra of the Maillard reaction between glucose and glycine catalyzed by Birnessite［20］
图5 美拉德反应产物与天然腐殖质的傅里叶变换红外光谱对比
（a）以葡萄糖与甘氨酸为底物的美拉德反应产物红外光谱；（b）不同类型天然土壤腐殖质的红外光谱［68］，其中A为红黄壤腐殖酸，B为深红壤腐殖酸。
Fig. 5 Comparison of Fourier Transform Infrared Spectra （FTIR） between Maillard reaction products and natural humic substances
（a） FTIR spectra of melanoidin-like polymers formed from glucose-glycine reactants； （b） FTIR spectra of various types of natural soil humic acids［68］， A is red yellow latosol humic acid， B is dark red latosol humic acid.
图6 土壤系统基于美拉德反应的矿物界面催化—类黑精生成—有机矿物复合体形成的有机碳三步稳定化路径
路径①~③代表传统腐殖化与有机碳固定，路径④~⑫代表美拉德反应介导的碳固定过程。①植物碳输入过程，主要以枯落物形式输入。②微生物对植被残体的分解过程，新碳形成老碳。③老碳在土壤中与矿物相结合形成稳定的有机—无机复合体。④新碳输入在土壤中迅速被微生物代谢转化［90-91］。⑤微生物分解有机碳和代谢过程产生糖类（d‑葡萄糖、d‑半乳糖、d‑核糖）、氨基酸（甘氨酸、l‑丙氨酸、l‑赖氨酸、l-异亮氨酸）等活性底物，为美拉德反应提供丰富的反应前体［92］。⑥部分代谢产物在土壤剖面中迁移并经由淋溶作用进入深层土壤，形成稳定的溶解有机碳库，成为深层微生物生长代谢和矿物界面催化反应的重要碳源［93］。⑦植物根系分泌物中直接富含葡萄糖、果糖、氨基酸等低分子有机物，这些底物无需微生物预处理即可直接进入非酶促美拉德反应，为根际及深层土壤提供即时反应基础［94］。⑧土壤中积累的老碳（主要来自植物残体）可被微生物缓慢分解利用，为土壤微生物提供有限但持续的碳源，支持其维持基础代谢活性。该过程体现了老碳矿化速率缓慢、时效性长的特征，且已被研究证实与根系分泌物与微生物交互行为密切相关，对碳循环和老碳激活具有调控作用［95］。⑨微生物分解老碳的过程中产生美拉德反应前体（还原性糖和氨基化合物）。⑩土壤中广泛存在的四价锰氧化物（如δ‑MnO2）通过氧化还原机制高效催化美拉德反应［18］。在该过程中，Mn4+作为氧化剂驱动还原糖与氨基化合物发生缩合和聚合反应，同时被还原为Mn2+［4］。随后，Mn2+可在分子氧或土著微生物的作用下再度氧化为Mn4+，形成具有循环活性的生物—无机催化体系［96］。土壤中多种细菌（如Pseudomonas putida， Leptothrix discophora）与真菌（如Phanerochaete chrysosporium）通过表达氧化酶或产生活性氧物种高效介导Mn2+→Mn4+的再氧化过程［97-98］，为类黑精的持续生成提供氧化驱动。⑪美拉德反应通过一系列的缩合、聚合反应生成具有高芳香性、高共轭度与分子复杂性的类黑精产物［99］。⑫生成的类黑精进一步与土壤中无机矿物（如黏土颗粒、氧化铁锰矿等）结合，形成有机—无机复合体从而被保存固定［93］。
Fig. 6 Three-step stabilization pathway of organic carbon in soil systems via the Maillard reaction： mineral interface catalysis， melanoidin formation， and organic-mineral complexation
Paths ①~③ represent traditional humification and organic carbon sequestration， while paths ④~⑫ represent Maillard reaction-mediated carbon sequestration processes. ① Plant carbon input， mainly in the form of litter. ② The decomposition of plant residues by microorganisms， where labile carbon is converted into recalcitrant carbon. ③ The combination of old carbon with minerals in the soil to form stable organic-inorganic complexes. ④ New carbon input is rapidly metabolized and transformed by microorganisms in the soil［90-91］. ⑤ Microbial decomposition of organic carbon and metabolic processes produce active substrates such as sugars （d-glucose， d-galactose， d-ribose） and amino acids （glycine， l-alanine， l-lysine， l-isoleucine）， providing abundant reaction precursors for the Maillard reaction［92］. ⑥ Some metabolic products migrate in the soil profile and enter the deep soil through leaching， forming a stable Dissolved Organic Carbon （DOC） pool， which becomes an important carbon source for deep soil microbial growth and metabolism and mineral interface catalytic reactions［93］. ⑦ Plant root exudates are directly rich in low-molecular-weight organic compounds such as glucose， fructose， and amino acids. These substrates can directly enter the non-enzymatic Maillard reaction without microbial pretreatment， providing an immediate reaction basis for the rhizosphere and deep soil［94］. ⑧ Old carbon accumulated in the soil （mainly from plant residues） can be slowly decomposed and utilized by microorganisms， providing a limited but continuous carbon source for soil microorganisms to maintain their basic metabolic activity. This process reflects the characteristics of slow mineralization rate and long-term effectiveness of old carbon， and has been confirmed to be closely related to the interaction between root exudates and microorganisms， playing a regulatory role in carbon cycling and old carbon activation［95］. ⑨ Maillard reaction precursors （reducing sugars and amino compounds） are produced during the microbial decomposition of old carbon. ⑩ The widely present tetravalent manganese oxides （such as δ-MnO2） in the soil efficiently catalyze the Maillard reaction through an oxidation-reduction mechanism［18］. In this process， Mn4+ acts as an oxidizing agent to drive the condensation and polymerization reactions of reducing sugars and amino compounds， while being reduced to Mn2+［4］. Subsequently， Mn2+ can be re-oxidized to Mn4+ under the action of molecular oxygen or indigenous microorganisms， forming a biologically-inorganic catalytic system with cyclic activity［96］. Various bacteria （such as Pseudomonas putida， Leptothrix discophora） and fungi （such as Phanerochaete chrysosporium） in the soil efficiently mediate the re-oxidation process of Mn2+→Mn4+ by expressing oxidases or producing reactive oxygen species［97-98］， providing the oxidative driving force for the continuous generation of melanoidins. ⑪ The Maillard reaction generates melanoidin products with high aromaticity， high conjugation， and molecular complexity through a series of condensation and polymerization reactions［99］. ⑫ The generated melanoidins further combine with inorganic minerals in the soil （such as clay particles， iron and manganese oxides， etc.）， forming organic-inorganic complexes and thus being preserved and fixed［93］.
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