塞罕坝地区优势造林树种叶片化学计量特征及其环境驱动

doi:10.17521/cjpe.2025.0161

植物生态学报 ›› 2026, Vol. 50 ›› Issue (2): 362-373.DOI: 10.17521/cjpe.2025.0161 cstr: 32100.14.cjpe.2025.0161

塞罕坝地区优势造林树种叶片化学计量特征及其环境驱动

田地¹(), 迟小龙¹, 石亮²^,^*(), 刘宵含¹, 赵常提¹, 吴梅¹, 张玉忠³, 高永亮⁴

¹ 北京林业大学, 林木资源高效生产全国重点实验室, 北京 100083
² 中国科学院地理科学与资源研究所, 北京 100101
³ 内蒙古自治区赤峰市克什克腾旗浩来呼热苏木人民政府林业工作站, 内蒙古赤峰 025349
⁴ 内蒙古克什克腾旗桦木沟林场, 内蒙古赤峰 025349

收稿日期:2025-05-06 接受日期:2025-11-29 出版日期:2026-02-28 发布日期:2026-04-01
通讯作者: *石亮 (shiliang0330@igsnrr.ac.cn)
基金资助:
国家重点研发计划(2022YFD2201600);中国科协青年人才托举工程(2021QNRC001)

Stoichiometric characteristics of dominant afforestation tree species and their environmental drivers in Saihanba region

TIAN Di¹(), CHI Xiao-Long¹, SHI Liang²^,^*(), LIU Xiao-Han¹, ZHAO Chang-Ti¹, WU Mei¹, ZHANG Yu-Zhong³, GAO Yong-Liang⁴

¹ State Key Laboratory of Efficient Production of Forest Resources, Beijing Forestry University, Beijing 100083, China
² Institute of Geographic Sciences and Resources, Chinese Academy of Sciences, Beijing 100101, China
³ Forestry Work Station of Haolaihure Sumu (Sub-district) People’s Government, Hexigten Banner, Chifeng, Nei Mongol 025349, China
⁴ Huamugou Forest Farm, Hexigten Banner, Chifeng, Nei Mongol 025349, China

Received:2025-05-06 Accepted:2025-11-29 Online:2026-02-28 Published:2026-04-01
Contact: *SHI Liang (shiliang0330@igsnrr.ac.cn)
Supported by:
National Key R&D Program of China(2022YFD2201600);Young Elite Scientists Sponsorship Program by the China Association for Science and Technology(2021QNRC001)

摘要/Abstract

摘要：

植物叶片化学计量特征与其光合作用、水分利用等生理生态功能密切相关。同一生境内不同树种的叶片化学计量特征反映出树木对环境养分资源的差异化利用策略。中国塞罕坝地区广布华北落叶松(Larix gmelinii var. principis-rupprechtii)和樟子松(Pinus sylvestris var. mongholica)人工林, 由于人工林传统经营中养分管理等方面的基础薄弱, 导致目前普遍存在生产力低、生态系统服务功能不强等问题。为了从树木养分利用的视角, 为半干旱区人工营林树种选择、生态系统提质增效提供理论依据, 该研究在塞罕坝地区设置野外调查固定样地, 采集了两种优势树种的叶片和土壤样品, 测定了其碳(C)、氮(N)和磷(P)含量及其化学计量比, 结合冗余分析和混合效应模型, 解析了两个树种叶片化学计量特征及其环境驱动。结果表明: 1)华北落叶松林土壤C、N和P含量高于樟子松林, 华北落叶松叶片N和P含量以及N含量相关养分计量比的种内变异均高于樟子松, 而叶片C含量低于樟子松; 2)在叶片化学计量特征的环境因子驱动方面, 土壤养分含量的影响力超过了大气水热因子, 土壤总碳含量解释了其变异的57.39%, 其次是年平均气温, 解释了其变异的15.77%; 3)随年平均气温升高, 华北落叶松叶片P含量下降, C:P和N:P显著上升, 而樟子松叶片化学计量特征无显著变化。可见, 塞罕坝地区的樟子松通过保持相对稳定的叶片N-P计量特征, 在适应胁迫环境时遵循“保守型”策略; 而华北落叶松则依赖较高的叶片N、P浓度和C:P、N:P, 呈现随环境条件而变化的“活跃型/获取型”策略。该研究明确了华北落叶松和樟子松叶片化学计量特征的差异, 为进一步开展养分调控树木生长实验, 解析两树种生理生态过程的养分调控机制提供了理论支撑。

关键词: 华北落叶松, 樟子松, 半干旱区, 叶片化学计量特征, 环境因子

Abstract:

Aims The stoichiometric characteristics of plant leaves are closely linked to their physiological and ecological functions, such as photosynthesis and water-use efficiency. Among tree species within the same habitat, variations in leaf stoichiometry reflect distinct strategies for utilizing environmental and nutritional resources. In China’s Saihanba region, extensive plantations of Larix gmeliniivar. principis-rupprechtii and Pinus sylvestrisvar. mongholica exhibit declining productivity and weakened ecosystem services, largely due to a lack of theoretical knowledge to inform traditional nutrient management. The objective of this study was to clarify the adaptive divergence and driving factors controlling the nutrient-use patterns of the dominant species used for afforestation projects and to use this knowledge to provide scientific guidance for tree species selection and ecosystem enhancement in semi-arid regions.

Methods We established permanent field plots in the Saihanba region and collected leaf and soil samples from both dominant species in these plots to measure their carbon (C), nitrogen (N), and phosphorus (P) content. We used Redundancy analysis (RDA) and mixed-effects models to identify key environmental drivers of variations in their leaf stoichiometry.

Important findings 1) Total soil C, N and P contents in the L. gmeliniivar. principis-rupprechtii forest were higher than those in the P. sylvestris var. mongholica forest. The total N and P contents in the leaves of L. gmeliniivar. principis-rupprechtii, as well as the intraspecific variation in N-related nutrient stoichiometric ratios, were all higher than those of P. sylvestris var. mongholica, while the leaf C content was lower than that of P. sylvestris var. mongholica; 2) In terms of the environmental factors driving the stoichiometric characteristics of leaves, the influence of soil nutrient content surpassed that of atmospheric variables. Total soil C content accounted for 57.39% of the variation, followed by the mean annual air temperature, which explained 15.77% of the variation; 3) With rising mean annual air temperature, leaf P content of L. gmeliniivar. principis-rupprechtiidecreased and leaf C:P and N:P ratios increased significantly. These findings suggest that P. sylvestris var. mongholica adopts a “conservative strategyˮ by maintaining relatively stable leaf N and P concentrations and constant N:P ratios as an adaptation to low-resource environments, while L. gmeliniivar. principis-rupprechtii employs an “active/acquisition strategyˮ characterized by higher N, P concentrations and flexible C:P and N:P ratios responsive to environmental shifts. This study elucidates the stoichiometric divergence between the two tree species, offering a theoretical foundation for future experiments on nutrient regulation and mechanistic exploration of their eco-physiological processes.

Key words: Larix gmelinii var. principis-rupprechtii, Pinus sylvestris var. mongholica, semi-arid region, leaf stoichiometric trait, environmental factor

田地, 迟小龙, 石亮, 刘宵含, 赵常提, 吴梅, 张玉忠, 高永亮. 塞罕坝地区优势造林树种叶片化学计量特征及其环境驱动. 植物生态学报, 2026, 50(2): 362-373. DOI: 10.17521/cjpe.2025.0161

TIAN Di, CHI Xiao-Long, SHI Liang, LIU Xiao-Han, ZHAO Chang-Ti, WU Mei, ZHANG Yu-Zhong, GAO Yong-Liang. Stoichiometric characteristics of dominant afforestation tree species and their environmental drivers in Saihanba region. Chinese Journal of Plant Ecology, 2026, 50(2): 362-373. DOI: 10.17521/cjpe.2025.0161

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2025.0161

https://www.plant-ecology.com/CN/Y2026/V50/I2/362

图/表 6

表1 塞罕坝野外调查样地的环境背景和林分特征(平均值±标准差)

Table 1 Environmental condition and forest stand information of the field experiment site in saihanba region (mean ± SD)

样地 Plot	树种 Tree species	纬度 Latitude (° N)	经度 Longitude (° E)	海拔 Altitude (m)	气温 Atmospheric temperature (℃)	大气湿度 Atmospheric humidity (%)	饱和水汽压差 Vapor pressure deficit (kPa)	土壤温度 Soil temperature (℃)	土壤湿度 Soil moisture (m³·m^-3)	林龄 Stand age (a)	平均胸径 Mean diameter at breast height (cm)	平均树高 Mean height (m)
1	LP	42.43	117.19	1 400	15.61 ± 4.30	71.76 ± 13.68	0.68 ± 0.45	14.01 ± 2.84	0.13 ± 0.03	40-45	17.88 ± 1.72	20.35 ± 1.50
2	LP	42.40	117.30	1 500	14.77 ± 4.49	71.81 ± 13.72	0.63 ± 0.43	11.84 ± 2.47	0.10 ± 0.02	40-45	18.43 ± 0.24	18.10 ± 1.69
3	LP	42.38	117.36	1 600	14.29 ± 4.30	76.71 ± 13.06	0.51 ± 0.39	12.82 ± 2.48	0.10 ± 0.02	45-50	15.73 ± 1.55	18.25 ± 1.73
4	PM	42.43	117.19	1 400	15.96 ± 4.09	71.54 ± 12.25	0.70 ± 0.46	13.24 ± 2.63	0.10 ± 0.04	45-50	15.56 ± 0.62	19.05 ± 1.63
5	PM	42.40	117.31	1 500	15.12 ± 4.52	70.52 ± 14.50	0.67 ± 0.46	10.23 ± 2.73	0.07 ± 0.01	40-45	16.04 ± 1.21	19.60 ± 0.81
6	PM	42.38	117.36	1 600	14.64 ± 4.41	72.26 ± 13.90	0.60 ± 0.42	10.58 ± 2.49	0.07 ± 0.01	45-50	15.90 ± 1.02	16.30 ± 2.06

图1 塞罕坝地区华北落叶松林和樟子松林土壤化学计量特征的差异。LP, 华北落叶松; PM, 樟子松。ns, p > 0.05; *, p < 0.05; **, p < 0.01; ****, p < 0.000 1。

Fig. 1 Comparative analysis of soil stoichiometric characteristics between larch and pine forests in the Saihanba region, China. LP, Larix gmelinii var. principis-rupprechtii; PM, Pinus sylvestris var. mongholica. Statistical significance is denoted as follows: ns, p > 0.05; *, p < 0.05; **, p < 0.01; ****, p < 0.000 1.

图2 塞罕坝地区华北落叶松和樟子松叶片化学计量特征对比。LP, 华北落叶松; PM, 樟子松。**, p < 0.01; ****, p < 0.000 1。

Fig. 2 Comparative analysis of leaf stoichiometric characteristics between larch and pine in the Saihanba region, China. LP, Larix gmelinii var. principis-rupprechtii; PM, Pinus sylvestris var. mongholica. Statistical significance is denoted as follows: **, p < 0.01; ****, p < 0.000 1.

图3 塞罕坝地区造林树种叶片化学计量特征影响因子的冗余分析(RDA)。蓝色虚线表示叶片的化学计量特征, 包括叶片碳含量(C)、叶片氮含量(N)、叶片磷含量(P)、叶片C:N、C:P和N:P; 黑色实线表示环境因子和土壤化学计量特征, 包括日均土壤水分含量(SM)、土壤全碳含量(STC)、土壤全氮含量(STN)、土壤全磷含量(STP)、年平均气温(T)、土壤温度(ST)和相对湿度(RH)。

Fig. 3 Redundancy analysis (RDA) of factors influencing leaf stoichiometric characteristics of afforestation tree species in the Saihanba region, China. Blue dashed arrows represent leaf stoichiometric characteristics, including leaf carbon content (C), nitrogen content (N), phosphorus content (P), C:N, C:P, and N:P. Black solid arrows represent environmental factors and soil stoichiometric characteristics, including annual mean soil moisture (SM), soil total carbon content (STC), soil total nitrogen content (STN), soil total phosphorus content (STP), annual mean air temperature (T), soil temperature (ST), and relative humidity (RH).

图4 塞罕坝地区华北落叶松和樟子松叶片化学计量特征对土壤全碳含量(STC)的响应。线性模型(粗黑线)和线性混合模型, 物种为随机变量(细彩色线)。线性混合效应模型边际或条件R2 (LMM: R2m、R2c)和线性模型R2 (LM)以及显著性(ns, p > 0.05; ***, p < 0.001)分别标注在数值的右上角。

Fig. 4 Univariate linear modeling of leaf stoichiometric characteristics and soil organic carbon content (STC). Both generalized (thick black lines) and linear mixed models including a random species intercept (thin colored lines) were fitted to the data. Corresponding model marginal or conditional R2 (LMM: R2m, R2c) and R2 (LM) and significance (ns, p > 0.05; ***, p < 0.001) are indicated on the top right corner of each value, respectively.

图5 塞罕坝地区华北落叶松和樟子松叶片化学计量学特征对年平均气温(T)的响应。拟合了线性模型(粗黑线)和线性混合模型, 物种为随机变量(细彩色线)。线性混合效应模型边际或条件R2 (LMM: R2m、R2c)和线性模型R2 (LM)以及显著性(ns, p > 0.05; *, p < 0.05; ***, p < 0.001)分别标注在数值的右上角。

Fig. 5 Univariate linear modeling of leaf stoichiometric characteristics and annual mean temperature (T). Both generalized (thick black lines) and linear mixed models including a random species intercept (thin colored lines) were fitted to the data. Corresponding model marginal or conditional R2 (LMM: R2m, R2c) and R2 (LM) and significance (ns, p > 0.05; *, p < 0.05; ***, p < 0.001) are indicated on the top right corner of each value, respectively.

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塞罕坝地区优势造林树种叶片化学计量特征及其环境驱动

Stoichiometric characteristics of dominant afforestation tree species and their environmental drivers in Saihanba region

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