中性糖在土壤中的来源与分布特征

doi:10.17521/cjpe.2018.0213

植物生态学报 ›› 2019, Vol. 43 ›› Issue (4): 284-295.DOI: 10.17521/cjpe.2018.0213

所属专题：微生物生态学

中性糖在土壤中的来源与分布特征

刘程竹^1,²,贾娟^1,²,戴国华¹,马田^1,²,冯晓娟^1,^2,^*()

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, 北京 100049

收稿日期:2018-08-27 修回日期:2019-04-18 出版日期:2019-04-20 发布日期:2019-08-29
通讯作者: 冯晓娟 ORCID:0000-0002-0443-0628
基金资助:
国家重点基础研究发展计划(973计划)(2015CB954201);国家自然科学基金(41773067);国家自然科学基金(41422304);中国科学院对外合作重点项目(151111KYSB20160014)

Origin and distribution of neutral sugars in soils

LIU Cheng-Zhu^1,²,JIA Juan^1,²,DAI Guo-Hua¹,MA Tian^1,²,FENG Xiao-Juan^1,^2,^*()

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

Received:2018-08-27 Revised:2019-04-18 Online:2019-04-20 Published:2019-08-29
Contact: FENG Xiao-Juan ORCID:0000-0002-0443-0628
Supported by:
Supported by the Chinese National Key Development Program for Basic Research(2015CB954201);the National Natural Science Foundation of China(41773067);the National Natural Science Foundation of China(41422304);the International Partnership Program of Chinese Academy of Sciences(151111KYSB20160014)

摘要/Abstract

摘要：

糖类(即碳水化合物)是土壤有机质的重要组成部分, 经生物化学降解形成不同结构的单糖。土壤中的中性单糖也叫中性糖, 主要包括木糖、核糖、阿拉伯糖、葡萄糖、半乳糖、甘露糖、岩藻糖和鼠李糖。其中, 植物来源的糖主要为五碳糖, 如木糖和阿拉伯糖; 微生物来源的糖主要包括半乳糖、甘露糖、岩藻糖、鼠李糖等六碳糖。研究中常利用六碳糖和五碳糖的比例指示微生物和植物对土壤有机碳的相对贡献。中性糖是微生物重要的碳源和能量来源, 在团聚体的形成过程中扮演着重要角色。该文整合了近30年土壤中性糖的研究进展, 对比了提取中性糖的常用方法, 分析了不同土地利用类型和不同土壤组分中中性糖的含量、来源和周转特征, 综述了影响中性糖含量和分布的主要环境因素。结果表明, 中性糖在耕地土壤中的绝对含量和相对含量均显著低于针叶林、阔叶林、草地和灌丛4种土地利用类型。(半乳糖+甘露糖)/(阿拉伯糖+木糖)(GM/AX)在不同土地利用间差异不显著, 而(鼠李糖+岩藻糖)/(阿拉伯糖+木糖)(RF/AX)则表明草地土壤中的微生物来源的中性糖含量高于针叶林和耕地。不同密度的土壤组分中, 轻质组分中中性糖的含量比重质组分高, 重质组分中微生物来源的中性糖较多; 就不同粒径(或团聚体)而言, 黏粒(或微团聚体)中微生物来源的中性糖含量更丰富。有关影响土壤中性糖含量和分布的因素的研究, 目前主要集中在人为活动(如耕种和放牧等), 而有关温度、降水等自然环境因素影响的研究较少。

关键词: 土壤, 中性糖, 来源, 分布特征, 影响因素

Abstract:

Carbohydrates are important components of soil organic matter, which can be decomposed to different types of monosaccharides. Neutral monosaccharides in the soil are also called neutral sugars, including xylose, ribose, arabinose, glucose, galactose, mannose, fucose and rhamnose. Among them, plant-derived sugars mainly include pentoses, such as xylose and arabinose, while microbial-derived sugars mainly consist of hexoses including galactose, mannose, fucose and rhamnose. Generally, the ratios of hexoses to pentoses are used to evaluate the contribution of microbial- versus plant-derived sugars. Neutral sugars are the main carbon and energy resources for soil microorganisms and play a vital role in aggregates formation. In this study, we review studies about neutral sugars in soils over the past 30 years and compare different methods for neutral sugar analysis. Furthermore, we compare the distribution patterns and turnover of soil neutral sugars across diverse land-use regimes, different soil density and particle size fractions and their influencing factors. The lowest neutral sugar content is found in arable soils compared with other four land-use types (coniferous forests, deciduous forests, shrublands and grasslands) in terms of absolute and relative contents. No significant difference is observed for the (galactose + mannose)/‍(arabinose + xylose)(GM/AX) ratios across the five land-use regimes. Nevertheless, the ratio of (rhamnose + fucose)/(arabinose + xylose)(RF/AX) indicates that microbially derived neutral sugars are more abundant in the soils of grasslands than coniferous forests or farmlands. The heavy fraction is characterized by an enrichment of microbial neutral sugars but a lower content of total neutral sugars compared to the light fraction. Concerning the distribution of neutral sugars across different soil size fractions (or aggregates), the microbial-derived neutral sugars are more abundant in the clay fraction (or microaggregates). As for the factors affecting neutral sugar content and distribution, many studies have focused on the human disturbances like agriculture and grazing, while the influence of environmental factors such as temperature, precipitation is poorly investigated.

Key words: soil, neutral sugars, origin, distribution, influencing factor

刘程竹, 贾娟, 戴国华, 马田, 冯晓娟. 中性糖在土壤中的来源与分布特征. 植物生态学报, 2019, 43(4): 284-295. DOI: 10.17521/cjpe.2018.0213

LIU Cheng-Zhu, JIA Juan, DAI Guo-Hua, MA Tian, FENG Xiao-Juan. Origin and distribution of neutral sugars in soils. Chinese Journal of Plant Ecology, 2019, 43(4): 284-295. DOI: 10.17521/cjpe.2018.0213

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2018.0213

https://www.plant-ecology.com/CN/Y2019/V43/I4/284

图/表 2

表1 土壤中性糖的提取及检测方法比较

Table 1 Comparison of extraction and detection methods of neutral sugars in soils

步骤 Procedure	方法 Method	优点 Advantage	缺点 Drawback	参考文献 Reference
提取 Extraction	硫酸 H₂SO₄		硫酸不易被除去 H₂SO₄ can not be removed easily	Tanaka et al., 1990
	盐酸 HCl		会水解一部分纤维素; 产率较低 The hydrolysis products include a few cellulosic neutral sugars; low yields	Uzaki & Ishiwatari, 1983
	三氟乙酸 TFA	产率高; 不会破坏单糖结构; 具有挥发性, 可通过旋转蒸发去除; 水解的多糖主要为半纤维素 High yields; Not destructive to monosaccharides; TFA is volatile and can be easily removed by evaporation; Hydrolysis products are mainly released from hemicellulose		Amelung et al., 1996
检测 Detection	GC-MS	精度、准确度、敏感性和效率较高 High accuracy, precision, sensibility and efficiency	需要衍生化 Derivatization is required	Amelung et al., 1996; Wang et al., 2017
	HPLC	无需衍生化; 纯化过程简单 No need for derivatization; Simple purification procedures	精度、准确度、敏感性较低 Low accuracy, precision, sensibility and efficiency	Hamada & Ono, 1984; Angers et al., 1988;Tanaka et al., 1990;
	HPAEC-PAD	无需衍生化; 应用范围广, 可同时分析糖醛酸和中性糖 No need for derivatization; Wide application and simultaneous analysis of uronic acid and neutral sugars	精度、准确度、敏感性较低 Low accuracy, precision, sensibility and efficiency	Bruggink et al., 2005; Zhang et al., 2012

图1 中性糖在不同土地利用类型表层土壤中的含量和来源(Nierop et al., 2001; Spielvogel et al., 2007; Eder et al., 2010; Rumpel et al., 2010; Zhao et al., 2014; Conti et al., 2016; Cui et al., 2016; Wang et al., 2016; Creme et al., 2017; Llorente et al., 2017; Evgrafova et al., 2018; Zhu et al., 2018)。A, 中性糖绝对含量。B, 中性糖相对含量。C, GM/AX ((半乳糖+甘露糖)/(阿拉伯糖+木糖))。D, RF/AX ((鼠李糖+岩藻糖)/(阿拉伯糖+木糖))。箱式图上方和下方的线段分别表示上四分位数和下四分位数,箱式图内部的横线表示数据的中位数,箱式图中在最上方或最下方的圆圈表示样本数据中的极端值。不同小写字母表示不同土地利用类型间差异显著(p < 0.05)。A, C, 阔叶林n = 8, 针叶林n = 25, 灌丛n = 4, 草地n = 8, 耕地n = 15。B, 阔叶林n = 8, 针叶林n = 22, 灌丛n = 4, 草地n = 8, 耕地n = 15。D, 阔叶林n = 3, 针叶林n = 19, 灌丛n = 4, 草地n = 8, 耕地n = 27。OC, 土壤有机碳。

Fig. 1 Content and distribution of neutral sugars across different land-use regimes in the top soils (Nierop et al., 2001; Spielvogel et al., 2007; Eder et al., 2010; Rumpel et al., 2010; Zhao et al., 2014; Conti et al., 2016; Cui et al., 2016; Wang et al., 2016; Creme et al., 2017; Llorente et al., 2017; Evgrafova et al., 2018; Zhu et al., 2018). A, Neutral sugar absolute content. B, Neutral sugar relative content. C, GM/AX ((galactose + mannose)/(arabinose + xylose)). D, RF/AX ((rhamnose + fucose)/(arabinose + xylose)). The upper and lower end of boxes denote the 0.25 and 0.75 percentiles, respectively. The solid bar in the box mark the median of each dataset. The circles indicate outliers of each dataset. Different lowercase letters indicate differences in various land-use regimes (p < 0.05). n = 8, 25, 4, 8, 15 (from deciduous, coniferous, shrub, grassland to crops in the A, C). n = 8, 22, 4, 8, 15 (from deciduous, coniferous, shrub, grassland to crops in the B). n = 3, 19, 4, 8, 27 (from deciduous, coniferous, shrub, grassland to crops in the D). OC, soil organic carbon.

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中性糖在土壤中的来源与分布特征

Origin and distribution of neutral sugars in soils

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