内蒙古典型草原土壤酶化学计量与限制性养分对放牧的响应

doi:10.17521/cjpe.2024.0113

植物生态学报 ›› 2025, Vol. 49 ›› Issue (1): 19-29.DOI: 10.17521/cjpe.2024.0113 cstr: 32100.14.cjpe.2024.0113

内蒙古典型草原土壤酶化学计量与限制性养分对放牧的响应

李天琦¹^,²(), 曹继容¹^,^*(), 柳小妮², 田思惠¹^,³, 兰波兰¹^,³, 邱颖¹, 薛建国¹, 张倩⁴, 褚建民⁴, 张淑敏¹, 黄建辉¹, 李凌浩¹, 王其兵¹

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, 国家植物园, 北京 100093
²甘肃农业大学草业学院, 兰州 730070
³中国科学院大学, 北京 100049
⁴中国林业科学院林业研究所, 北京 100091

收稿日期:2024-04-17 接受日期:2024-12-10 出版日期:2025-01-20 发布日期:2025-03-08
通讯作者: * (jrcao@ibcas.ac.cn)
作者简介:李天琦: ORCID: 0009-0001-5868-9133
基金资助:
中国科学院战略性先导科技专项(A类)(XDA26020102);国家自然科学基金(42077056);国家自然科学基金(42477519);国家自然科学基金(41977046);国家重点研发计划(2023YFF404105)

Response of soil enzyme stoichiometry to grazing and identification of soil limiting nutrients in typical steppe of Nei Mongol, China

LI Tian-Qi¹^,²(), CAO Ji-Rong¹^,^*(), LIU Xiao-Ni², TIAN Si-Hui¹^,³, LAN Bo-Lan¹^,³, QIU Ying¹, XUE Jian-Guo¹, ZHANG Qian⁴, CHU Jian-Min⁴, ZHANG Shu-Min¹, HUANG Jian-Hui¹, LI Ling-Hao¹, WANG Qi-Bing¹

¹State Key Laboratory of Vegetation and Environmental Change, Institute of Botany, Chines Academy of Sciences, China National Botanical Garden, Beijing 100093, China
²College of Pratacultural Science, Gansu Agricultural University, Lanzhou 730070, China
³University of Chinese Academy of Sciences, Beijing 100049, China
⁴Research Institute of Forestry, Chinese Academy of Forestry, Beijing 100091, China

Received:2024-04-17 Accepted:2024-12-10 Online:2025-01-20 Published:2025-03-08
Supported by:
Strategic Priority Research Program of the Chinese Academy of Sciences(XDA26020102);National Natural Science Foundation of China(42077056);National Natural Science Foundation of China(42477519);National Natural Science Foundation of China(41977046);National Key R&D Program of China(2023YFF404105)

摘要/Abstract

摘要：

土壤胞外酶介导有机物分解, 维持生态系统内养分循环, 其活性及化学计量比是反映土壤养分可利用性、解析微生物资源限制的重要指标。然而放牧, 特别是过度放牧引起的内蒙古典型草原退化对土壤胞外酶化学计量和养分限制特征的影响尚不清楚。该研究以内蒙古典型草原为研究对象, 基于放牧强度梯度实验, 分析了不同放牧强度下土壤胞外酶活性及其化学计量比的变化, 并通过矢量模型解析了土壤养分限制特征。结果表明: (1)研究区域土壤水解酶活性在0-300 nmol·g^-1·h^-1范围内, 与全球土壤酶库中的相应数值相比偏低。放牧强度显著影响α-葡萄糖苷酶、纤维素水解酶、木糖苷酶、纤维素二糖苷酶、β-1,4-N-乙酰氨基葡萄糖苷酶、亮氨酸氨基肽酶、酸性磷酸酶活性。随放牧强度的增加, 酶活性动态呈现“双峰曲线”格局, 分别在推荐放牧(RG)和中度放牧(MG)下出现活性峰值。(2)标准主轴回归分析表明, 碳循环酶活性和氮循环酶活性、磷循环酶活性三者之间存在显著的线性关系。土壤酶碳:氮:磷化学计量比约为1:2.3:1.3, 偏离了全球1:1:1的结果。(3)基于土壤酶化学计量比的矢量模型分析表明内蒙古典型草原放牧草地受到氮磷共同限制, 磷限制更强烈, 且随着放牧强度的增强, 磷限制加剧。

关键词: 草地退化, 放牧, 土壤酶活性, 养分限制

Abstract:

Aims Soil extracellular enzymes are crucial for soil organic matter decomposition and nutrient cycling. Soil enzyme activity and stoichiometry can provide insights into microbial resource limitations and soil nutrient availability. This study investigated the effects of grazing, particularly overgrazing that often leads to grassland degradation, on soil enzyme activity and stoichiometry, and identifies nutrient limitations in temperate grasslands.

Methods We conducted grazing experiments with varying stock rates in a typical steppe of Nei Mongol, and investigated the changes in the activities and stoichiometric ratios of soil extracellular enzymes. Enzyme activities related to carbon (C), nitrogen (N), and phosphorus (P) cycling were analyzed, and a vector model was applied to determine soil nutrient limitations under different grazing intensities.

Important findings 1) Soil hydrolase activities in the studied grassland ranged from 0 to 300 nmol·g^-1·h^-1, which is relatively low compared with the global averages. Grazing intensity significantly impacted the activities of soil enzymes, including α-glucosidase, cellulose hydrolysis, xylosidase, β-d-cellubiosidase, β-1,4-N-acetylamino-glucosidase, glycosaminidase, leucine aminopeptidase, and acid phosphatase. The enzyme activities peaked under moderate grazing and recommended grazing. 2) The Standardized Major Axis (SMA) regression analysis revealed strong linear relationships between the enzyme activities associated with C, N, and P cycling. The soil enzyme C:N:P stoichiometric ratio was 1:2.3:1.3, deviating from the global average 1:1:1. 3) The vector model based on soil enzyme stoichiometry indicated that the grasslands were co-limited by N and P, with P limitation becoming more pronounced as grazing intensity increased in Nei Mongol.

Key words: grassland degradation, grazing, soil enzyme activity, nutrient limitation

李天琦, 曹继容, 柳小妮, 田思惠, 兰波兰, 邱颖, 薛建国, 张倩, 褚建民, 张淑敏, 黄建辉, 李凌浩, 王其兵. 内蒙古典型草原土壤酶化学计量与限制性养分对放牧的响应. 植物生态学报, 2025, 49(1): 19-29. DOI: 10.17521/cjpe.2024.0113

LI Tian-Qi, CAO Ji-Rong, LIU Xiao-Ni, TIAN Si-Hui, LAN Bo-Lan, QIU Ying, XUE Jian-Guo, ZHANG Qian, CHU Jian-Min, ZHANG Shu-Min, HUANG Jian-Hui, LI Ling-Hao, WANG Qi-Bing. Response of soil enzyme stoichiometry to grazing and identification of soil limiting nutrients in typical steppe of Nei Mongol, China. Chinese Journal of Plant Ecology, 2025, 49(1): 19-29. DOI: 10.17521/cjpe.2024.0113

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https://www.plant-ecology.com/CN/Y2025/V49/I1/19

图/表 5

表1 放牧强度对土壤胞外酶活性的影响(平均值±标准误, nmol·g-1·h-1, n = 20)

Table 1 Effect of grazing intensity on soil extracellular enzyme activities (mean ± SE, nmol·g-1·h-1, n = 20)

土壤酶 Soil enzyme	放牧强度 Grazing intensity
土壤酶 Soil enzyme	CK	RG	LG	MG	HG
AG	8.73 ± 0.61^a	8.13 ± 0.40^ab	7.43 ± 0.41^ab	7.75 ± 0.62^ab	6.78 ± 0.43^b
BG	11.03 ± 0.89^a	11.58 ± 1.06^a	9.47 ± 0.42^a	8.98 ± 1.00^a	9.05 ± 0.61^a
CBH	1.70 ± 0.25^b	0.79 ± 0.36^c	1.87 ± 0.33^ab	2.01 ± 0.23^ab	2.67 ± 0.23^a
XS	8.33 ± 0.73^ab	8.82 ± 0.47^a	6.70 ± 0.64^bc	7.55 ± 0.85^ab	4.91 ± 0.58^c
CB	5.47 ± 0.46^a	5.40 ± 0.29^a	4.60 ± 0.42^ab	5.00 ± 0.52^a	3.48 ± 0.30^b
NAG	12.40 ± 0.65^ab	14.52 ± 2.53^a	11.21 ± 0.39^ab	10.76 ± 0.69^ab	9.40 ± 0.51^b
LAP	257.56 ± 14.78^a	254.98 ± 14.28^a	217.66 ± 13.10^ab	229.61 ± 17.56^ab	198.45 ± 7.94^b
AP	26.36 ± 3.29^ab	28.56 ± 2.80^a	17.93 ± 3.10^b	22.21 ± 3.47^ab	8.84 ± 1.37^c

图1 不同放牧强度下土壤碳氮磷循环相关酶活性的变化(平均值±标准误, n = 20)。不同小写字母表示不同放牧强度间差异显著(p < 0.05)。AP, 酸性磷酸酶; BG, β-葡萄糖苷酶; LAP, 亮氨酸氨基肽酶; NAG, β-1,4-N-乙酰氨基葡萄糖苷酶。CK, 对照; HG, 重度放牧; LG, 轻度放牧; MG, 中度放牧; RG, 推荐放牧。

Fig. 1 Soil extracellular enzyme activities associated with carbon (C), nitrogen (N) and phosphorus (P) acquisition under different grazing intensities (mean ± SE, n = 20). Significant differences among grazing intensities are indicated by different lowercase letters (p < 0.05). AP, acid phosphatase; BG, β-1,4-glucosidase; LAP, leucine aminopeptidase; NAG, β-N-acetyl glucosaminidas. CK, control; HG, heavy grazing; LG, light grazing; MG, moderate grazing; RG, recommended grazing.

图2 不同放牧强度下土壤胞外酶化学计量比(平均值±标准误, n = 20)。不同小写字母表示不同放牧强度间差异显著(p < 0.05)。C, 碳; N, 氮; P, 磷。CK, 对照; HG, 重度放牧; LG, 轻度放牧; MG, 中度放牧; RG, 推荐放牧。

Fig. 2 Soil extracellular enzyme stoichiometric ratios under different grazing intensities (mean ± SE, n = 20). Significant differences among grazing intensities are denoted by different lowercase letters (p < 0.05). C, carbon; N, nitrogen; P, phosphorus. CK, control; HG, heavy grazing; LG, light grazing; MG, moderate grazing; RG, recommended grazing.

图3 酸性磷酸酶(AP)、β-葡萄糖苷酶(BG)、亮氨酸氨基肽酶(LAP)、β-1,4-N-乙酰氨基葡萄糖苷酶(NAG)活性关系的标准主轴回归分析(n = 100)。CK, 对照; HG, 重度放牧; LG, 轻度放牧; MG, 中度放牧; RG, 推荐放牧。

Fig. 3 Standard Major Axis (SMA) regression analyses illustrating the relationships between the activities of acid phosphatase (AP), β-1,4-glucosidase (BG), leucine aminopeptidase (LAP) and β-N-acetyl glucosaminidase (NAG) (n = 100). CK, control; HG, heavy grazing; LG, light grazing; MG, moderate grazing; RG, recommended grazing.

图4 不同放牧强度下土壤酶化学计量的矢量模型分析。A, 矢量长度随放牧强度的变化(平均值±标准误)。B, 矢量角度随放牧强度的变化，其中虚线为45°阈值线(平均值±标准误)。C, 土壤酶碳氮化学计量比与碳磷化学计量比的关系, 虚线为1:1阈值线。D, 矢量长度与矢量角度的线性拟合以甄别限制性养分。不同小写字母表示不同放牧强度间差异显著(p < 0.05)。CK, 对照; HG, 重度放牧; LG, 轻度放牧; MG, 中度放牧; RG, 推荐放牧。

Fig. 4 Vector model analysis based on soil extracellular enzyme stoichiometric ratios under different grazing intensities. A, Variation of vector length with grazing intensity (mean ± SE). B, Variation of vector angle with grazing intensity, the dotted 45° threshold line distinguishes nitrogen (N) and phosphorus (P) limitation (mean ± SE). C, Relationship between soil enzyme carbon (C):N and C:P stoichiometric ratios, with the dashed line indicating the 1:1 threshold. D, Linear regression analysis of vector length and angle, highlighting patterns of nutrient limitation. Different lowercase letters denote significant differences among grazing intensities (p < 0.05). CK, control; HG, heavy grazing; LG, light grazing; MG, moderate grazing; RG, recommended grazing.

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内蒙古典型草原土壤酶化学计量与限制性养分对放牧的响应

Response of soil enzyme stoichiometry to grazing and identification of soil limiting nutrients in typical steppe of Nei Mongol, China

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