暖温带森林土壤原生生物功能类群对凋落物与根系资源变化的差异性响应

doi:10.17520/biods.2025441

生物多样性 ›› 2026, Vol. 34 ›› Issue (6): 25441. DOI: 10.17520/biods.2025441 cstr: 32101.14.biods.2025441

• 研究报告: 微生物多样性 • 上一篇下一篇

暖温带森林土壤原生生物功能类群对凋落物与根系资源变化的差异性响应

胡莎莎¹^,², 李志鹏²^,^*()(), 张卫信⁴(), 姚海凤²^,³(), 马磊⁴, 张晓⁵(), 刘晓静⁶, 龚鑫²(), 孙新¹^,²(), 傅声雷⁴^,^*()()

¹ 福建农林大学生命科学学院, 福州 350002
² 中国科学院城市环境研究所城市环境与健康重点实验室/福建省流域生态重点实验室, 福建厦门 361021
³ 中国科学院大学, 北京 100049
⁴ 河南大学, 郑州 450046
⁵ 西北农林科技大学, 陕西杨凌 712100
⁶ 河南内乡宝天曼国家级自然保护区管理局, 河南内乡 474350

收稿日期:2025-11-05 接受日期:2026-04-22 出版日期:2026-06-20 发布日期:2026-07-30
通讯作者: *共同通讯作者 E-mail: zpli@iue.ac.cn; fsl@vip.henu.edu.cn
基金资助:
国家自然科学基金(32301430);国家自然科学基金(32361143523);国家重点研发计划(2023YFF1304601)

Divergent responses of protist functional groups to litter and root resource changes in a warm-temperate forest

Shasha Hu¹^,², Zhipeng Li²^,^*()(), Weixin Zhang⁴(), Haifeng Yao²^,³(), Lei Ma⁴, Xiao Zhang⁵(), Xiaojing Liu⁶, Xin Gong²(), Xin Sun¹^,²(), Shenglei Fu⁴^,^*()()

¹ College of Life Sciences, Fujian Agriculture and Forestry University, Fuzhou 350002, China
² Key Laboratory of Urban Environment and Health/Fujian Key Laboratory of Watershed Ecology, Institute of Urban Environment, Chinese Academy of Sciences, Xiamen, Fujian 361021, China
³ University of Chinese Academy of Sciences, Beijing 100049, China
⁴ Henan University, Zhengzhou 450046, China
⁵ Northwest A&F University, Yangling, Shaanxi 712100, China
⁶ Baotianman National Nature Reserve Administration, Neixiang, Henan 474350, China

Received:2025-11-05 Accepted:2026-04-22 Online:2026-06-20 Published:2026-07-30
Contact: *Co-authors for correspondence. E-mail: zpli@iue.ac.cn; fsl@vip.henu.edu.cn
Supported by:
National Natural Science Foundation of China(32301430);National Natural Science Foundation of China(32361143523);National Key Research and Development Program of China(2023YFF1304601)

1. 附录.pdf(777KB)

摘要/Abstract

摘要： 植物光合产物主要以地上凋落物和根系输入的形式进入土壤, 为地下食物网提供物质与能量。原生生物作为土壤生物多样性的核心组分, 在微生物调控方面发挥重要作用。然而, 凋落物与根系输入对土壤原生生物多样性与群落结构的影响及机制并不清楚。本研究基于宝天曼森林生态系统国家野外科学观测研究站连续6.5年的资源控制实验(移除凋落物与灌木和阻断乔木根系), 应用环境DNA宏条形码技术分析了食细菌者、食真核生物者、杂食者和食碎屑者4类原生生物功能群对植物资源变化的响应。结果表明, 不同原生生物功能群对植物资源变化的响应存在显著差异。总体上, 移除凋落物显著降低了各原生生物功能群的丰富度, 其降幅约为33.7%–47.7% (P < 0.05)。移除凋落物通过降低细菌和真菌丰富度对各原生生物功能类群丰富度产生“自下而上”的影响, 其中, 食细菌者与食真核生物者主要与细菌丰富度下降有关, 食碎屑者与真菌丰富度下降有关, 而杂食者类群则受到细菌和真菌丰富度下降的共同驱动。阻断根系的作用相对较弱, 且阻断灌木根系和乔木根系的效应在不同原生生物功能类群中存在明显差异。阻断乔木根系降低了多个类群的丰富度, 并通过增加土壤含水量间接提高了食真核生物者和食碎屑者丰富度。总体而言, 凋落物与根系资源通过不同路径共同调控土壤原生生物丰富度, 不同原生生物功能类群对两类植物资源的差异化响应, 与其受到土壤性质与微生物资源的差异化调控密切相关。本研究通过揭示原生生物对植物资源输入的响应差异模式, 强调原生生物在连接地上–地下过程、维持土壤微食物网稳定性中的关键作用, 为深入理解土壤生态系统中原生生物的功能机制提供依据。

关键词: 资源调控实验, 原生生物功能类群, 环境DNA, 土壤食物网, 暖温带落叶阔叶林

Abstract

Aims: Plant photosynthetic products enter soil primarily through litter and root inputs, providing material and energy for belowground food webs. As a key component of soil biodiversity, protists play a crucial role in microbial regulation. However, the effects and mechanisms of litter and root inputs on the structure and diversity of soil protist communities remain unclear.

Methods: Based on a six-and-a-half-year resource manipulation experiment including litter removal and root deprivation at the Baotianman Forest Ecosystem National Observation and Research Station, eDNA metabarcoding was used to analyze the responses of four protist functional groups (bacterivores, eukaryvores, omnivores, and detritivores) to changes in plant resource inputs.

Results: Litter removal significantly reduced the richness of all protist functional groups, with declines ranging from 33.7% to 47.7% (P < 0.05). Litter removal exerted strong bottom-up negative effects on richness of protists by reducing bacterial and fungal richness. Specifically, bacterivores and eukaryvores were primarily associated with declines in bacterial richness, detritivores were mainly linked to reductions in fungal richness, whereas omnivores were jointly driven by decreases in both bacterial and fungal richness. The effects of root deprivation were relatively weaker, and the impacts of the two root resource types differed markedly among protist functional groups. Tree root deprivation exerted direct negative effects on the richness of multiple protist groups (except detritivores), while indirectly exerting positive effects on eukaryvores and detritivores by increasing soil moisture. By comparison, shrub root deprivation mainly promoted fungal richness, thereby indirectly exerting positive effects on omnivores and detritivores. Litter and root resources jointly regulate soil protist diversity through distinct pathways, and the differential responses of protist functional groups to these two plant resource inputs are closely linked to their contrasting effects on soil properties and microbial resources.

Conclusions: These findings reveal the diverse responses of protists to changes in plant resource inputs and highlight their crucial roles in linking aboveground and belowground processes, maintaining micro-food web stability, thereby providing new insights into the functional mechanisms of protists in soil ecosystems.

Key words: resource manipulation experiment, functional groups of protists, environmental DNA, soil foodweb, warm temperate deciduous broad leaved forest

胡莎莎, 李志鹏, 张卫信, 姚海凤, 马磊, 张晓, 刘晓静, 龚鑫, 孙新, 傅声雷 (2026) 暖温带森林土壤原生生物功能类群对凋落物与根系资源变化的差异性响应. 生物多样性, 34, 25441. DOI: 10.17520/biods.2025441.

Shasha Hu, Zhipeng Li, Weixin Zhang, Haifeng Yao, Lei Ma, Xiao Zhang, Xiaojing Liu, Xin Gong, Xin Sun, Shenglei Fu (2026) Divergent responses of protist functional groups to litter and root resource changes in a warm-temperate forest. Biodiversity Science, 34, 25441. DOI: 10.17520/biods.2025441.

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链接本文: https://www.biodiversity-science.net/CN/10.17520/biods.2025441

https://www.biodiversity-science.net/CN/Y2026/V34/I6/25441

图/表 9

图1 移除凋落物、阻断乔木根系和阻断灌木根系的复合处理示意图

Fig. 1 Diagram of combined treatment of litter removal, tree root deprivation and shrub root deprivation

图2 实验样地设置空间示意图

Fig. 2 Spatial schematic diagram of the experimental plot design

表1 基于双因素方差分析(ANOVA)的移除凋落物和阻断根系对土壤理化性质和微生物资源丰富度的影响。加粗字体代表差异显著(P < 0.05)。

Table 1 Effects of litter removal and root deprivation on soil physicochemical properties and microbial resource richness based on two-way analysis of variance (ANOVA). Bold font indicates significant differences (P < 0.05). TP, Total phosphorus; TC, Total carbon; TN, Total nitrogen; TS, Total sulfur.

土壤理化性质及微生物资源丰富度 Soil properties and microbial resources richness	移除凋落物 Litter removal (df = 1)		阻断根系 Root deprivation (df = 2)		交互作用 Interaction (df = 2)
	F	P	F	P	F	P
含水率 Moisture content (%)	3.42	0.08	4.42	< 0.05	0.68	0.52
酸碱度 pH	0.22	0.65	0.58	0.57	2.37	0.12
总碳 TC (g/kg)	6.29	< 0.05	1.88	0.18	1.75	0.20
总氮 TN (g/kg)	21.00	< 0.001	2.51	0.11	1.85	0.18
总磷 TP (g/kg)	5.76	< 0.05	0.47	0.63	0.17	0.84
总硫 TS (g/kg)	4.72	< 0.05	0.0001	0.10	0.31	0.74
真菌丰富度 Fungal richness	20.22	< 0.001	2.74	0.09	0.55	0.59
细菌丰富度 Bacterial richness	27.75	< 0.001	3.55	0.05	3.76	< 0.05

图3 移除凋落物和阻断根系对土壤理化性质和微生物资源丰富度的影响。不同字母表示不同根系处理间存在显著差异; 星号表示不同凋落物处理间存在显著差异。*** P < 0.001, ** P < 0.01, * P < 0.05。

Fig. 3 Effects of litter removal and root deprivation on soil physicochemical properties and microbial resource richness. Different letters indicate significant differences among root treatments; asterisks indicate significant differences between litter treatments. *** P < 0.001, ** P < 0.01, * P < 0.05. TP, Total phosphorus; TC, Total carbon; TN, Total nitrogen; TS, Total sulfur.

表2 基于双因素方差分析(ANOVA)的移除凋落物和阻断根系对不同原生生物功能类群丰富度的影响。加粗字体代表差异显著(P < 0.05)。

Table 2 Effects of litter removal and root deprivation on the richness of different protists functional groups based on two-way analysis of variance (ANOVA). Bold font indicates significant differences (P < 0.05).

原生生物功能类群 Functional groups of protists	移除凋落物 Litter removal (df = 1)		阻断根系 Root deprivation (df = 2)		交互作用 Interaction (df = 2)
原生生物功能类群 Functional groups of protists	F	P	F	P	F	P
食细菌者 Bacterivores	44.30	< 0.001	6.20	< 0.01	0.38	0.70
食真核生物者 Eukaryvores	30.59	< 0.001	7.40	< 0.01	4.12	< 0.05
杂食者 Omnivores	54.52	< 0.001	4.76	< 0.05	0.27	0.77
食碎屑者 Detritivores	9.75	< 0.01	2.73	0.09	4.16	< 0.05

图4 移除凋落物和阻断根系对食细菌者(A)、食真核生物者(B)、杂食者(C)和食碎屑者(D)丰富度的影响。不同字母表示不同根系处理间存在显著差异; 星号表示不同凋落物处理间存在显著差异。*** P < 0.001,** P < 0.01, * P < 0.05。

Fig. 4 Effects of litter removal and root deprivation on the richness of bacterivore (A), eukaryvore (B), omnivore (C), and detritivore (D) protists. Different letters indicate significant differences among root treatments. Asterisks denote significant differences between litter treatments. *** P < 0.001, ** P < 0.01, * P < 0.05.

图5 移除凋落物和阻断根系处理下食细菌者、食真核生物者、杂食者和食碎屑者群落组成的主坐标分析(PCoA)。颜色表示凋落物处理, 点形状和椭圆线型表示根系处理。椭圆表示95%的置信区间; R2表示移除凋落物和阻断根系对群落组成变异的解释度。

Fig. 5 Principal coordinate analysis (PCoA) of community composition of bacterivores, eukaryvores, omnivores, and detritivores protist under litter removal and root deprivation. Colors indicate litter treatments, while point shapes and ellipse line types represent root treatments. Ellipses denote 95% confidence intervals. R2 indicates the proportion of variation in community composition explained by litter removal and root deprivation.

图6 分段结构方程模型(PSEM)展示移除凋落物、阻断根系、土壤理化性质和微生物资源对土壤原生生物功能类群丰富度的直接和间接影响。红色箭头表示正相关性, 蓝色箭头表示负相关性; 实线表示显著(P < 0.05), 虚线表示不显著(P > 0.05)。*** P < 0.001, ** P < 0.01, * P < 0.05。线条旁的数值是标准化路径系数; 线条的粗细表示路径系数的显著程度; 作用越强线条越粗。R2表示模型中各变量对土壤理化性质、微生物资源以及原生生物功能类群丰富度变化的解释度。TC: 总碳; TN: 总氮; TS: 总硫; TP: 总磷。

Fig. 6 Piecewise structural equation models (PSEM) showing the direct and indirect effects of litter removal, root deprivation, and soil physicochemical properties and microbial resources on the richness of soil protist functional groups. Red arrows indicate positive correlations, and blue arrows indicate negative correlations. Solid lines represent significant effects (P < 0.05), whereas dashed lines represent non-significant effects (P > 0.05). *** P < 0.001, ** P < 0.01, * P < 0.05. Values adjacent to arrows indicate standardized path coefficients. Line thickness indicates the significance of path coefficients; thicker lines represent stronger effects. R² indicates the proportion of variance in soil physicochemical properties, microbial resources and protist functional group richness explained by the model variables. TC, Total carbon; TN, Total nitrogen; TS, Total sulfur; TP, Total phosphorus.

表3 分段结构方程模型(PSEM)中移除凋落物及阻断根系对原生生物功能类群丰富度显著的直接、间接及总效应

Table 3 Significant direct, indirect, and total effects of litter removal and root deprivation on the richness of protist functional groups in the piecewise structural equation models (PSEM)

原生生物功能类群 Functional groups of protists	效应 Effects	移除凋落物 Litter removal	阻断灌木根系 Shrub root deprivation	阻断乔木根系 Tree root deprivation
食细菌者 Bacterivores	直接效应 Direct effects	0	0	-0.35
	间接效应 Indirect effects	-0.61	0	0
	总效应 Total effects	-0.61	0	-0.35
食真核生物者 Eukaryvores	直接效应 Direct effects	0	0	-0.33
	间接效应 Indirect effects	-0.61	0	0.27
	总效应 Total effects	-0.61	0	-0.06
杂食者 Omnivores	直接效应 Direct effects	0	-0.32	-0.55
	间接效应 Indirect effects	-0.77	0.13	0
	总效应 Total effects	-0.77	-0.19	-0.55
食碎屑者 Detritivores	直接效应 Direct effects	0	0	0
	间接效应 Indirect effects	-0.49	0.23	0
	总效应 Total effects	-0.49	0.23	0

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暖温带森林土壤原生生物功能类群对凋落物与根系资源变化的差异性响应

Divergent responses of protist functional groups to litter and root resource changes in a warm-temperate forest

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