植物生态学报 ›› 2026, Vol. 50 ›› Issue (4): 859-869.DOI: 10.17521/cjpe.2025.0119 cstr: 32100.14.cjpe.2025.0119
张安宁1, 肖亚宁2, 赵霞2, 张淼1, 崔瀚文2, 陈书燕1,*(
), 安黎哲1,3,*(
)
收稿日期:2025-04-02
接受日期:2025-09-09
出版日期:2026-04-20
发布日期:2026-06-25
通讯作者:
*陈书燕, chenshy@lzu.edu.cn;基金资助:
ZHANG An-Ning1, XIAO Ya-Ning2, ZHAO Xia2, ZHANG Miao1, CUI Han-Wen2, CHEN Shu-Yan1,*(
), AN Li-Zhe1,3,*(
)
Received:2025-04-02
Accepted:2025-09-09
Online:2026-04-20
Published:2026-06-25
Contact:
*CHEN Shu-Yan, chenshy@lzu.edu.cn;Supported by:摘要: 线虫是地下生物群落的重要组成部分, 在维持生态系统结构与功能中起着至关重要的作用。全球环境变化, 如灌丛化和氮沉降, 已成为生物群落结构和功能变化的重要驱动因素。不同环境因子会共同影响生物多样性, 而灌丛化和氮沉降间的交互作用对土壤线虫功能多样性的影响尚不清晰。该研究在青藏高原东缘开展了灌木移除和氮添加的控制实验, 使用核密度N维超体积法计算线虫功能多样性, 探究线虫对灌木、氮沉降及其交互作用的响应。结果表明: (1)灌木显著增加了线虫物种多样性并改变了其群落组成, 但该效应受氮添加影响较小。(2)灌木显著增加了线虫功能多样性, 而氮添加将削弱灌木对功能多样性的促进作用。(3)灌木改变了不同食性线虫多度占比, 显著增加了植食性、食细菌性和食真菌性线虫占比, 但显著降低了捕食性线虫占比。此外, 氮添加加剧了灌木对低营养级线虫的促进作用和对高营养级线虫的抑制作用。(4)相关性和冗余分析(RDA)表明, 植物生物量和土壤水分含量是影响线虫群落组成和功能性状的关键驱动因子。在灌丛化和大气氮沉降背景下, 线虫功能多样性将更加相似。
张安宁, 肖亚宁, 赵霞, 张淼, 崔瀚文, 陈书燕, 安黎哲. 青藏高原灌丛化与氮添加对土壤线虫群落及功能性状的交互作用. 植物生态学报, 2026, 50(4): 859-869. DOI: 10.17521/cjpe.2025.0119
ZHANG An-Ning, XIAO Ya-Ning, ZHAO Xia, ZHANG Miao, CUI Han-Wen, CHEN Shu-Yan, AN Li-Zhe. Interactions between shrub encroachment and nitrogen addition on nematode community and functional traits on Qingzang Plateau. Chinese Journal of Plant Ecology, 2026, 50(4): 859-869. DOI: 10.17521/cjpe.2025.0119
图1 灌木移除、氮添加及其交互作用对线虫物种多样性的影响。不同小写字母表示差异显著(p < 0.05)。
Fig. 1 Effects of shrub removal, nitrogen (N) addition, and their interaction on the nematode taxonomic diversity. Different lowercase letters indicate significant difference (p < 0.05).
图2 灌木移除、氮添加及其交互作用对土壤线虫群落组成的影响。CC, 灌木移除及不添加氮; CN, 灌木移除及添加氮; SC, 灌木存在及不添加氮; SN, 灌木存在及添加氮。NMDS, 非度量多维尺度分析。
Fig. 2 Effects of shrub removal, nitrogen (N) addition, and their interaction on the nematode community composition. CC, shrub removal and no N addition; CN, shrub removal and N addition; SC, with shrub and no N addition; SN, with shrub and N addition. NMDS, non-metric multidimensional scaling analysis.
图3 灌木移除、氮添加及其交互作用对线虫属相对丰度影响的堆积图。CC, 灌木移除及不添加氮; CN, 灌木移除及添加氮; SC, 灌木存在及不添加氮; SN, 灌木存在及添加氮。
Fig. 3 Stacked bar of the effects of shrub removal, nitrogen addition, and their interaction on the relative abundance of nematode genera. CC, shrub removal and no nitrogen addition; CN, shrub removal and nitrogen addition; SC, with shrub and no nitrogen addition; SN, with shrub and nitrogen addition.
| 线虫属 Nematode | 对照 Control | 氮添加 Nitrogen addition | p | ||||
|---|---|---|---|---|---|---|---|
| 灌木存在 With shrub | 灌木移除 Shrub removal | 灌木存在 With shrub | 灌木移除 Shrub removal | 灌木效应 Shrub | 氮效应 Nitrogen | 交互作用 Shrub × nitrogen | |
| 拟丽突属 Acrobeloides | 12.44 ± 3.69a | 23.43 ± 4.99a | 15.64 ± 4.16a | 2.40 ± 1.17b | 0.075 | <0.001 | <0.001 |
| 中杆属 Mesorhabditis | 10.31 ± 4.30ab | 1.94 ± 1.16b | 10.94 ± 24.45a | 3.08 ± 1.82ab | 0.012 | 0.464 | 0.635 |
| 原矛属 Prodorylaimium | 4.82 ± 2.28b | 1.82 ± 1.04b | 0.91 ± 0.53b | 15.21 ± 4.37a | 0.946 | 0.076 | 0.002 |
| 垫刃属 Tylenchus | 7.95 ± 3.44a | 2.81 ± 1.68a | 9.27 ± 3.63a | 2.50 ± 1.36a | 0.167 | 0.864 | 0.746 |
| 真矛属 Eudorylaimus | 3.47 ± 1.91ab | 0.84 ± 0.58b | 3.51 ± 1.92ab | 9.45 ± 4.45a | 0.034 | <0.001 | 0.004 |
| 类滑刃属 Aphelenchoides | 4.26 ± 2.87a | 4.30 ± 2.57a | 3.66 ± 2.31a | 1.58 ± 1.10a | 0.986 | 0.083 | 0.279 |
| 滑刃属 Aphelenchus | 7.04 ± 1.86a | 0.46 ± 0.26b | 9.86 ± 2.21a | 0.22 ± 0.13b | <0.001 | 0.258 | 0.138 |
| 盘旋属 Rotylenchus | 8.72 ± 3.55a | 1.38 ± 0.82b | 2.98 ± 1.62ab | 2.57 ± 1.43ab | 0.002 | 0.318 | 0.045 |
| 单齿属 Mononchus | 1.26 ± 0.76b | 1.27 ± 0.76b | 2.42 ± 1.35ab | 10.08 ± 3.73a | 0.998 | <0.001 | 0.086 |
| 盆咽属 Panagrolaimus | 2.77 ± 1.52a | 2.66 ± 1.36a | 1.80 ± 0.97a | 4.30 ± 2.21a | 0.937 | 0.341 | 0.246 |
| 异头叶属 Pseudacrobeles | 5.80 ± 1.36a | 0.57 ± 0.29b | 7.61 ± 1.56a | 0.11 ± 0.07c | <0.001 | 0.010 | 0.007 |
| 锉齿属 Mylonchulus | 1.81 ± 1.16a | 5.50 ± 2.83a | 2.27 ± 1.25a | 2.36 ± 1.30a | 0.097 | 0.184 | 0.219 |
| 钵盘属 Mylodiscus | 2.12 ± 1.19ab | 7.22 ± 2.95a | 1.13 ± 0.69b | 1.22 ± 0.73b | 0.035 | 0.003 | 0.167 |
| 大矛属 Enchodelus | 0.33 ± 0.22b | 0.41 ± 0.28b | 0.41 ± 0.28b | 10.51 ± 3.12a | 0.733 | <0.001 | <0.001 |
| 绕线属 Plectus | 5.01 ± 1.00a | 0.12 ± 0.07b | 6.42 ± 1.14a | 0.07 ± 0.04b | <0.001 | 0.373 | 0.231 |
表1 灌木移除、氮添加及其交互作用对前15个线虫属丰度的影响(平均值±标准误)
Table 1 Effects of shrub removal, nitrogen addition, and their interaction on the abundance of first 15 nematode genera (mean ± SE)
| 线虫属 Nematode | 对照 Control | 氮添加 Nitrogen addition | p | ||||
|---|---|---|---|---|---|---|---|
| 灌木存在 With shrub | 灌木移除 Shrub removal | 灌木存在 With shrub | 灌木移除 Shrub removal | 灌木效应 Shrub | 氮效应 Nitrogen | 交互作用 Shrub × nitrogen | |
| 拟丽突属 Acrobeloides | 12.44 ± 3.69a | 23.43 ± 4.99a | 15.64 ± 4.16a | 2.40 ± 1.17b | 0.075 | <0.001 | <0.001 |
| 中杆属 Mesorhabditis | 10.31 ± 4.30ab | 1.94 ± 1.16b | 10.94 ± 24.45a | 3.08 ± 1.82ab | 0.012 | 0.464 | 0.635 |
| 原矛属 Prodorylaimium | 4.82 ± 2.28b | 1.82 ± 1.04b | 0.91 ± 0.53b | 15.21 ± 4.37a | 0.946 | 0.076 | 0.002 |
| 垫刃属 Tylenchus | 7.95 ± 3.44a | 2.81 ± 1.68a | 9.27 ± 3.63a | 2.50 ± 1.36a | 0.167 | 0.864 | 0.746 |
| 真矛属 Eudorylaimus | 3.47 ± 1.91ab | 0.84 ± 0.58b | 3.51 ± 1.92ab | 9.45 ± 4.45a | 0.034 | <0.001 | 0.004 |
| 类滑刃属 Aphelenchoides | 4.26 ± 2.87a | 4.30 ± 2.57a | 3.66 ± 2.31a | 1.58 ± 1.10a | 0.986 | 0.083 | 0.279 |
| 滑刃属 Aphelenchus | 7.04 ± 1.86a | 0.46 ± 0.26b | 9.86 ± 2.21a | 0.22 ± 0.13b | <0.001 | 0.258 | 0.138 |
| 盘旋属 Rotylenchus | 8.72 ± 3.55a | 1.38 ± 0.82b | 2.98 ± 1.62ab | 2.57 ± 1.43ab | 0.002 | 0.318 | 0.045 |
| 单齿属 Mononchus | 1.26 ± 0.76b | 1.27 ± 0.76b | 2.42 ± 1.35ab | 10.08 ± 3.73a | 0.998 | <0.001 | 0.086 |
| 盆咽属 Panagrolaimus | 2.77 ± 1.52a | 2.66 ± 1.36a | 1.80 ± 0.97a | 4.30 ± 2.21a | 0.937 | 0.341 | 0.246 |
| 异头叶属 Pseudacrobeles | 5.80 ± 1.36a | 0.57 ± 0.29b | 7.61 ± 1.56a | 0.11 ± 0.07c | <0.001 | 0.010 | 0.007 |
| 锉齿属 Mylonchulus | 1.81 ± 1.16a | 5.50 ± 2.83a | 2.27 ± 1.25a | 2.36 ± 1.30a | 0.097 | 0.184 | 0.219 |
| 钵盘属 Mylodiscus | 2.12 ± 1.19ab | 7.22 ± 2.95a | 1.13 ± 0.69b | 1.22 ± 0.73b | 0.035 | 0.003 | 0.167 |
| 大矛属 Enchodelus | 0.33 ± 0.22b | 0.41 ± 0.28b | 0.41 ± 0.28b | 10.51 ± 3.12a | 0.733 | <0.001 | <0.001 |
| 绕线属 Plectus | 5.01 ± 1.00a | 0.12 ± 0.07b | 6.42 ± 1.14a | 0.07 ± 0.04b | <0.001 | 0.373 | 0.231 |
图4 灌木移除、氮添加及其交互作用对土壤线虫功能多样性的影响。不同小写字母表示差异显著(p < 0.05)。
Fig. 4 Effects of shrub removal, nitrogen (N) addition, and their interaction on the nematode functional diversity. Different lowercase letters indicate significant difference (p < 0.05).
图5 灌木移除、氮添加及其交互作用对土壤线虫群落功能性状的影响。A, 线虫C-P值的群落加权平均值。B, 线虫生物量的群落加权平均值。C, 食细菌性线虫占比。D, 食真菌性线虫占比。E, 植食性线虫占比。F, 捕食性线虫占比。G, 杂食性线虫占比。占比为多度占比。不同小写字母表示差异显著(p < 0.05)。
Fig. 5 Effects of shrub removal, nitrogen (N) addition, and their interaction on the functional traits of nematode communities. A, Community weighted mean of nematode C-P value. B, Community weighted mean of nematode biomass. C, Bacterivore nematode proportion. D, Fungivore nematode proportion. E, Herbivore nematode proportion. F, Predator nematode proportion. G, Omnivore nematode proportion. Proportion refers to abundance proportion. Different lowercase letters indicate significant difference (p < 0.05).
| 线虫指标 Nematode indicator | 土壤 pH | 电导率 Electrical conductivity | 含水量 Soil moisture | 土壤总磷含量 Total phosphorus content | 土壤硝态氮含量 Ammonium nitrogen content | 土壤铵态氮含量 Nitrate nitrogen content | 植物生物量 Plant biomass | 植物丰富度 Plant richness |
|---|---|---|---|---|---|---|---|---|
| 物种多样性 Taxonomic diversity | -0.096 | 0.168 | 0.083 | -0.535* | 0.043 | 0.025 | -0.253 | -0.032 |
| 功能多样性 Functional diversity | -0.001 | 0.090 | 0.184 | -0.342 | 0.072 | -0.145 | -0.171 | -0.020 |
| C-P值 C-P value | 0.139 | -0.125 | 0.412 | 0.108 | -0.498* | 0.089 | 0.350 | 0.275 |
| 线虫生物量 Biomass | 0.233 | 0.195 | 0.554* | -0.458* | -0.394 | 0.060 | -0.055 | 0.094 |
| 食细菌性占比 Bacterivore proportion | -0.068 | 0.199 | -0.341 | -0.044 | 0.462* | -0.131 | -0.301 | -0.154 |
| 食真菌性占比 Fungivore proportion | -0.047 | 0.072 | -0.175 | -0.321 | 0.216 | 0.063 | 0.486* | -0.356 |
| 植食性占比 Herbivore proportion | -0.136 | 0.136 | -0.372 | -0.134 | 0.168 | 0.013 | -0.278 | -0.227 |
| 杂食性占比 Omnivore proportion | -0.103 | -0.170 | 0.165 | 0.091 | -0.226 | 0.206 | 0.397 | 0.078 |
| 捕食性占比 Predator proportion | 0.269 | -0.191 | 0.546* | 0.224 | -0.522* | -0.076 | 0.415 | 0.427 |
表2 土壤线虫多样性及功能性状与环境因子间的相关关系
Table 2 Correlation between the diversity and functional traits of soil nematodes and environmental factors
| 线虫指标 Nematode indicator | 土壤 pH | 电导率 Electrical conductivity | 含水量 Soil moisture | 土壤总磷含量 Total phosphorus content | 土壤硝态氮含量 Ammonium nitrogen content | 土壤铵态氮含量 Nitrate nitrogen content | 植物生物量 Plant biomass | 植物丰富度 Plant richness |
|---|---|---|---|---|---|---|---|---|
| 物种多样性 Taxonomic diversity | -0.096 | 0.168 | 0.083 | -0.535* | 0.043 | 0.025 | -0.253 | -0.032 |
| 功能多样性 Functional diversity | -0.001 | 0.090 | 0.184 | -0.342 | 0.072 | -0.145 | -0.171 | -0.020 |
| C-P值 C-P value | 0.139 | -0.125 | 0.412 | 0.108 | -0.498* | 0.089 | 0.350 | 0.275 |
| 线虫生物量 Biomass | 0.233 | 0.195 | 0.554* | -0.458* | -0.394 | 0.060 | -0.055 | 0.094 |
| 食细菌性占比 Bacterivore proportion | -0.068 | 0.199 | -0.341 | -0.044 | 0.462* | -0.131 | -0.301 | -0.154 |
| 食真菌性占比 Fungivore proportion | -0.047 | 0.072 | -0.175 | -0.321 | 0.216 | 0.063 | 0.486* | -0.356 |
| 植食性占比 Herbivore proportion | -0.136 | 0.136 | -0.372 | -0.134 | 0.168 | 0.013 | -0.278 | -0.227 |
| 杂食性占比 Omnivore proportion | -0.103 | -0.170 | 0.165 | 0.091 | -0.226 | 0.206 | 0.397 | 0.078 |
| 捕食性占比 Predator proportion | 0.269 | -0.191 | 0.546* | 0.224 | -0.522* | -0.076 | 0.415 | 0.427 |
图6 土壤线虫群落分布与环境因子关系的冗余分析(RDA)二维排序图。红线表示线虫属, 蓝线表示环境因子。EC, 电导率; NH+4-N, 土壤铵态氮含量; NO-3-N, 土壤硝态氮含量; PB, 植物生物量; pH, 土壤pH; PR, 植物丰富度; SM, 土壤含水量; TP, 土壤总磷含量。Acrobeloides, 拟丽突属; Alaimus, 无咽属; Allodorylaimus, 异矛线属; Anaplectus, 似绕线属; Aphelenchoides, 类滑刃属; Aphelenchus, 滑刃属; Aporcelaimus, 孔咽属; Cephalobus, 头叶属; Enchodelus, 大矛属; Filenchus, 丝垫刃属; Iotonchus, 基齿属; Labronema, 厚唇属; Miconchus, 等齿属; Mononchus, 单齿属; Mylodiscus, 钵盘属; Mylonchulus, 锉齿属; Plectus, 绕线属; Prodorylaimium, 原矛线属; Protorhabditis, 原杆属; Pseudacrobeles, 异头叶属; Rotylenchus, 盘旋属; Panagrolaimus, 盆咽属; Paratimminema, 库氏线属; Tylenchorhynchus, 矮化属; Tylenchus, 垫刃属; Tylopharynx, 瘤咽属。部分属名位于原点附近, 被遮盖, 有Coomansus, 库曼属; Criconemoides, 轮属; Dorylaimoides, 拟矛线属; Dorylaimus, 矛线属; Epidorylaimus, 附矛线属; Eudorylaimus, 真矛属; Helicotylenchus, 螺旋属; Laimydorus, 咽矛线属; Mesodorylaimus, 中矛线属; Mesorhabditis, 中杆属; Mononchoides, 拟独齿属。
Fig. 6 Redundancy analysis (RDA) of two-dimensional diagram of the relationship between soil nematode community distribution and environment factors. Red lines represent nematode genera, and blue lines represent environmental factors. EC, electrical conductivity; NH+4-N, ammonium nitrogen content; NO-3-N, nitrate nitrogen content; PB, plant biomass; pH, soil pH; PR, plant richness; SM, soil moisture; TP, total phosphorus content.
| 变量 Variable | df | F | p |
|---|---|---|---|
| 土壤含水量 Soil moisture | 1 | 2.505 | 0.010 |
| 植物生物量 Plant biomass | 1 | 2.180 | 0.027 |
| 土壤总磷含量 Soil total phosphorus content | 1 | 1.622 | 0.092 |
| 电导率 Electrical conductivity | 1 | 1.211 | 0.247 |
| 土壤pH Soil pH | 1 | 0.891 | 0.547 |
| 土壤硝态氮含量 Soil ammonium nitrogen content | 1 | 1.313 | 0.191 |
| 土壤铵态氮含量 Soil nitrate nitrogen content | 1 | 1.254 | 0.223 |
| 植物丰富度 Plant richness | 1 | 0.838 | 0.581 |
| 残差 Residual | 11 |
表3 环境因子对土壤线虫群落组成影响的冗余分析
Table 3 Redundancy analysis of the effect of environment factors on the nematode community composition
| 变量 Variable | df | F | p |
|---|---|---|---|
| 土壤含水量 Soil moisture | 1 | 2.505 | 0.010 |
| 植物生物量 Plant biomass | 1 | 2.180 | 0.027 |
| 土壤总磷含量 Soil total phosphorus content | 1 | 1.622 | 0.092 |
| 电导率 Electrical conductivity | 1 | 1.211 | 0.247 |
| 土壤pH Soil pH | 1 | 0.891 | 0.547 |
| 土壤硝态氮含量 Soil ammonium nitrogen content | 1 | 1.313 | 0.191 |
| 土壤铵态氮含量 Soil nitrate nitrogen content | 1 | 1.254 | 0.223 |
| 植物丰富度 Plant richness | 1 | 0.838 | 0.581 |
| 残差 Residual | 11 |
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