华北不同林龄人工林植物功能性状与碳氮化学计量

doi:10.17521/cjpe.2025.0183

摘要/Abstract

摘要：

为明确不同树种人工林植物功能性状与碳氮化学计量在林龄梯度下的变化规律及响应机制, 该研究以不同林龄毛白杨(Populus tomentosa)、侧柏(Platycladus orientalis)和国槐(Styphnolobium japonicum)人工林为研究对象, 测定并探究其不同器官功能性状、碳氮化学计量和土壤理化性质的变化及相互关系。研究发现: (1)随着林龄的增加, 3个树种土壤养分、黏粒和粉粒含量均显著增加, 毛白杨土壤全磷含量、侧柏土壤pH减小; (2) 3个树种比叶面积均随林龄增加而显著增加, 比叶质量显著减小。根系碳含量(平均值为36.25%)低于枝条(45.52%)与叶片(44.83%), 叶片氮含量(1.98%)高于根系(1.30%)与枝条(0.64%), 不同器官养分含量与碳氮比随林龄增加的变化因树种而异; (3)毛白杨各器官功能性状间存在协同一致性, 而侧柏叶片与根性状独立演化, 形成策略性解耦; (4)土壤含水量的增加与土壤质地的改善是引起植物性状变化的重要原因; (5)土壤全氮含量与叶片和根系碳含量呈显著负相关关系, 冗余分析表明土壤电导率是影响植物功能性状的显著因子; (6)随机森林分析表明, 影响植物叶片、枝条与根系碳氮比的显著因素包括土壤全磷和黏粒含量、比叶面积、叶干物质量与植物养分含量。研究结果揭示了不同树种在林龄梯度下植物功能性状与器官碳氮分配的差异, 丰富了植物经济谱理论框架。

关键词: 林龄, 功能性状, 植物经济谱, 碳氮比

Abstract:

Aims Clarify the variation patterns and corresponding response mechanisms of plant functional traits and carbon-nitrogen stoichiometry in artificial plantations of different tree species under the age gradient.

Methods In this study, Populus tomentosa, Platycladus orientalis and Styphnolobium japonicum at different ages were selected as the research objects to determine and explore the changes and relationships of different organ functional traits, carbon-nitrogen stoichiometry and soil physicochemical properties.

Important findings (1) With the increase of forest age, the content of soil nutrients, clay and silt of different tree plantations increased significantly. While soil total phosphorus content of Populus tomentosa plantations, and soil pH value of Platycladus orientalis plantations decreased. (2) The specific leaf area of different tree species increased significantly with the increase of forest age, while the specific leaf mass decreased significantly. The root carbon content (average 36.25%) was lower than that of branches (45.52%) and leaves (44.83%), and the leaf nitrogen content (1.98%) was higher than that of roots (1.30%) and branches (0.64%). With the increase of forest age, the nutrient content and carbon-nitrogen ratio of different organs varied with tree species. (3) The functional traits of Populus tomentosa plantations of different organs were coordinated, while the functional traits of leaves and roots of Platycladus orientalisplantations evolved independently, forming a decoupling strategy. (4) The increase of soil water content and the improvement of soil texture are important reasons for the changes in plant traits. (5) The soil total nitrogen content was significantly negatively correlated with the carbon content in leaves and roots. Redundancy analysis showed that soil electrical conductivity significantly affected plant functional traits. (6) Random forest analysis showed that the significant factors affecting the carbon-nitrogen ratio of leaves, branches and roots included soil total phosphorus and clay contents, specific leaf area, leaf dry matter mass and plant nutrient content. The research results reveal the differences in plant functional traits and organ carbon and nitrogen distribution among different tree species under the forest age gradient, enriching the theoretical framework of the plant economic spectrum.

Key words: forest age, functional trait, plant economic spectrum, carbon to nitrogen ratio

陈雅轩, 韩雨音, 陈艳梅, 刘倩愿. 华北不同林龄人工林植物功能性状与碳氮化学计量. 植物生态学报, 2026, 50(2): 318-333. DOI: 10.17521/cjpe.2025.0183

CHEN Ya-Xuan, HAN Yu-Yin, CHEN Yan-Mei, LIU Qian-Yuan. Plant functional traits and carbon-nitrogen stoichiometry of plantations at different forest ages in North China. Chinese Journal of Plant Ecology, 2026, 50(2): 318-333. DOI: 10.17521/cjpe.2025.0183

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2025.0183

https://www.plant-ecology.com/CN/Y2026/V50/I2/318

图/表 9

表1 各树种人工林样地基本概况

Table 1 Basic information of the artificial plantations sample plots of each tree species

树种 Species	林龄 Forest age (a)	简称 Abbreviation	经纬度 Longitude and latitude	海拔 Altitude (m)	郁闭度 Canopy density (%)	平均胸径 Average diameter at breast height (cm)	平均树高 Average height (m)
毛白杨 Populus tomentosa	10	XY	37.49° N, 114.38° E	169	15	6.5	4.6
	16	ZY	37.48° N, 114.38° E	174	20	10.5	7.2
	30	LY	37.50° N, 114.39° E	157	25	22.3	13.4
侧柏 Platycladus orientalis	12	XB	37.49° N, 114.39° E	165	10	2.4	2.5
	19	ZB	37.50° N, 114.39° E	185	60	6.8	5.5
	36	LB	37.52° N, 114.39° E	246	80	18.1	7.2
国槐 Styphnolobium japonicum	9	XH	37.45° N, 114.41° E	99	10	4.0	3.7
	13	ZH	37.45° N, 114.41° E	47	40	10.1	9.5
	16	LH	37.51° N, 114.39° E	199	65	12.5	7.1

表2 植物功能性状属性

Table 2 Attributes of plant functional traits

功能性状 Functional trait	缩写 Abbreviation	单位 Unit	计算公式 Calculation formula
叶片含水量 Leaf water content	LWC	%	LWC = (LFM-LDM)/LFM
比叶面积 Specific leaf area	SLA	cm²∙g^-1	SLA = LA/LDM
比叶质量 Leaf mass per area	LMA	g∙cm^-2	LMA = LDM/LA
叶干物质含量 Leaf dry matter content	LDMC	g∙g^-1	LDMC = LDM/LFM
叶片氮含量 Leaf nitrogen content	LNC	%
叶片碳含量 Leaf carbon content	LCC	%
叶片碳氮比 Leaf carbon to nitrogen ratio	L_C:N		L_C:N =LCC/LNC
枝条氮含量 Branch nitrogen content	BNC	%
枝条碳含量 Branch carbon content	BCC	%
枝条碳氮比 Branch carbon to nitrogen ratio	B_C:N		B_C:N =BCC/BNC
根系生物量 Root biomass	RB	kg∙m^-3	RB = RDM/SV
根平均直径 Root average diameter	RD	mm
根长密度 Root length density	RLD	km∙m^-3	RLD = RL/SV
根表面积密度 Root surface area density	RAD	m²∙m^-3	RAD = RA/SV
比根长 Specific root length	SRL	m∙g^-1	SRL = RL/RDM
根比表面积 Specific root area	SRA	m²∙g^-1	SRA = RA/RDM
根组织密度 Root tissue density	RTD	g∙cm^-3	RTD = RDM/RV
根氮含量 Root nitrogen content	RNC	%
根碳含量 Root carbon content	RCC	%
根碳氮比 Root carbon to nitrogen content	R_C:N		R_C:N = RCC/RNC

表3 华北不同林龄人工林土壤理化性质变化

Table 4 Changes in soil physicochemical properties at different forest ages in north China

图1 华北不同林龄人工林植物功能性状变化。不同小写字母表示同一树种在不同林龄间的差异显著(p < 0.05)。

Fig. 1 Changes in functional traits of plantations at different forest ages in north China. Different lowercase letters indicate the significant differences of the same species at different forest ages (p < 0.05).

图2 不同林龄人工林植物碳、氮含量及化学计量变化。不同小写字母表示同一树种在不同林龄间的差异显著(p < 0.05)。

Fig. 2 Changes in carbon (C), nitrogen (N) content and stoichiometry of plantations at different ages. Different lowercase letters indicate the significant differences of the same species at different forest ages (p < 0.05).

图3 叶片性状(A)与根系性状(B)的主成分(PC)分析。PO, 侧柏人工林; PT, 毛白杨人工林; SJ, 国槐人工林。植物功能性状缩写含义见表2。***, p < 0.001。

Fig. 3 Principal component (PC) analysis of leaf traits (A) and root traits (B). PO, Platycladus orientalis plantations; PT, Populus tomentosa plantations; SJ, Styphnolobium japonicum plantations. Abbreviations of plant functional traits are shown in Table 2. ***, p < 0.001.

图4 土壤理化性质、植物功能性状与化学计量相关性分析。C:N, 土壤碳氮比; C:P, 土壤碳磷比; Clay, 土壤黏粒含量; EC, 土壤电导率; N:P, 土壤氮磷比; pH, 土壤pH; Sand, 土壤砂粒含量; Silt, 土壤粉粒含量; TN, 土壤全氮含量; TOC, 土壤总有机碳含量; TP, 土壤全磷含量; WC, 土壤含水量。植物功能性状缩写含义见表2。*, p < 0.05; **, p < 0.01; ***, p < 0.001。

Fig. 4 Correlations among soil physicochemical properties, plant functional traits, and stoichiometry. C:N, soil carbon to nitrogen ratio; C:P, soil carbon to phosphorus ratio; Clay, soil clay content; EC, soil electrical conductivity; N:P, soil nitrogen to phosphorus ratio; pH, soil pH; Sand, soil sand content; Silt, soil silt content; TN, soil total nitrogen content; TOC, soil total organic carbon content; TP, soil total phosphorus content; WC, soil water content. Abbreviations of plant functional traits are shown in Table 2. *, p < 0.05; **, p < 0.01; ***, p < 0.001.

图5 叶片、根系功能性状与土壤理化性质的冗余分析(A)与解释率(B)。C:N, 土壤碳氮比; C:P, 土壤碳磷比; Clay, 土壤黏粒含量; EC, 土壤电导率; N:P, 土壤氮磷比; pH, 土壤pH; Sand, 土壤砂粒含量; Silt, 土壤粉粒含量; TN, 土壤全氮含量; TOC, 土壤总有机碳含量; TP, 土壤全磷含量; WC, 土壤含水量。植物功能性状缩写含义见表2。***, p < 0.001。

Fig. 5 Redundancy analysis (A) and explanation rate (B) of leaf and root functional traits, and soil physical and chemical properties. C:N, soil carbon to nitrogen ratio; C:P, soil carbon to phosphorus ratio; Clay, soil clay content; EC, soil electrical conductivity; N:P, soil nitrogen to phosphorus ratio; pH, soil pH; Sand, soil sand content; Silt, soil silt content; TN, soil total nitrogen content; TOC, soil total organic carbon content; TP, soil total phosphorus content; WC, soil water content. Abbreviations of plant functional traits are shown in Table 2. ***, p < 0.001.

图6 影响叶片(A)、枝条(B)与根系(C)碳氮比的变量相对重要性排序。C:N, 土壤碳氮比; C:P, 土壤碳磷比; Clay, 土壤黏粒含量; EC, 土壤电导率; N:P, 土壤氮磷比; pH, 土壤pH; Sand, 土壤砂粒含量; Silt, 土壤粉粒含量; TN, 土壤全氮含量; TOC, 土壤总有机碳含量; TP, 土壤全磷含量; WC, 土壤含水量。植物功能性状缩写含义见表2。Var explained为方差解释率。*, p < 0.05; **, p < 0.01。

Fig. 6 Relative importance of indicators affecting carbon to nitrogen ratio of leaves (A), branches (B), and roots (C). C:N, soil carbon to nitrogen ratio; C:P, soil carbon to phosphorus ratio; Clay, soil clay content; EC, soil electrical conductivity; N:P, soil nitrogen to phosphorus ratio; pH, soil pH; Sand, soil sand content; Silt, soil silt content; TN, soil total nitrogen content; TOC, soil total organic carbon content; TP, soil total phosphorus content; WC, soil water content. Abbreviations of plant functional traits are shown in Table 2. Var, variance; X.IncMSE, Variable x’s increase in mean squared error. *, p < 0.05; **, p < 0.01.

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