器官年龄在油松叶、枝、根化学计量特征变异以及协调中的重要作用

doi:10.17521/cjpe.2025.0097

植物生态学报 ›› 2026, Vol. 50 ›› Issue (2): 306-317.DOI: 10.17521/cjpe.2025.0097 cstr: 32100.14.cjpe.2025.0097

器官年龄在油松叶、枝、根化学计量特征变异以及协调中的重要作用

于江珊¹, 许浩²^,³^,⁴, 郭永忠²^,³^,⁴, 侯继华¹^,^*()

¹ 北京林业大学生态与自然保护学院, 北京 100083
² 宁夏农林科学院林业与草地生态研究所, 银川 750002
³ 宁夏防沙治沙与水土保持重点实验室, 银川 750002
⁴ 宁夏生态修复与多功能林业综合研究中心, 银川 750002

收稿日期:2025-03-19 接受日期:2025-05-13 出版日期:2026-02-28 发布日期:2026-04-01
通讯作者: *侯继华 (houjihua@bjfu.edu.cn)
基金资助:
国家自然科学基金(31872683)

Important role of organ age in variation and coordination of stoichiometry in Pinus tabuliformis leaves, twigs and roots

YU Jiang-Shan¹, XU Hao²^,³^,⁴, GUO Yong-Zhong²^,³^,⁴, HOU Ji-Hua¹^,^*()

¹ School of Ecology and Nature Conservation, Beijing Forestry University, Beijing 100083, China
² Institute of Forestry and Grassland Ecology, Ningxia Academy of Agricultural and Forestry Sciences, Yinchuan 750002, China
³ Ningxia Key Laboratory of Desertification Control and Soil and Water Conservation, Yinchuan 750002, China
⁴ Research Center for Ecological Restoration and Multi-Functional Forestry of Ningxia, Yinchuan 750002, China

Received:2025-03-19 Accepted:2025-05-13 Online:2026-02-28 Published:2026-04-01
Contact: *HOU Ji-Hua (houjihua@bjfu.edu.cn)
Supported by:
National Natural Science Foundation of China(31872683)

摘要/Abstract

摘要：

油松(Pinus tabuliformis)为中国温带针叶林的建群种和关键物种, 具有较强的环境适应能力。研究油松当年生器官(新生器官)和多年生器官(≥1年生, 成熟器官)的叶、枝、根的化学计量特征及其相互关系, 对于深入理解植物种内变异和资源分配策略具有重要意义。该研究选取了中国9个地区的天然油松林, 测定了81株油松的当年生叶、多年生叶、当年生枝、多年生枝、吸收根和运输根的碳(C)、氮(N)和磷(P)含量。结果表明: (1)当年生器官的N和P含量高于多年生器官, 而多年生器官的C:N、C:P和N:P比值则高于当年生器官。不同年龄组之间, 各器官的化学计量特征表现出不同程度的种内变异, 且器官年龄是决定变异的主要因素。(2)标准主轴回归分析显示, 相较于当年生叶, 多年生叶的P含量更高; 相较于多年生枝和运输根, 当年生枝和吸收根的C、N、P含量更高。(3)油松吸收根的C、N、P含量显著高于当年生叶和当年生枝。(4)冗余分析表明, 影响当年生叶和多年生叶化学计量特征的主要环境因子分别为年平均气温和土壤P含量; 对当年生枝和多年生枝化学计量特征影响最大的环境因子分别为年降水量和土壤P含量; 而影响吸收根和运输根化学计量特征的主要环境因子分别为土壤含水率和土壤N含量。该研究揭示了油松不同年龄组叶、枝、根的资源分配策略及其环境适应性。该研究结果表明, 在研究常绿树种化学计量特征时, 应充分考虑各器官的年龄差异, 这对阐明其养分分配和环境适应策略至关重要。

关键词: 种内变异, 油松, 叶, 枝, 根, 年龄, 化学计量特征

Abstract:

Aims As a dominant and keystone species in temperate coniferous forests of China, Pinus tabuliformis exhibits strong environmental adaptability. Investigating the stoichiometric characteristics of leaves, twigs and roots in both current-year (newly developed) and perennial (≥1 year old, mature) organs, as well as their relationships, is essential for understanding intraspecific variation and plant resource allocation strategies.

Methods This study was conducted in natural P. tabuliformis forests across nine regions in China. A total of 81 individual trees were sampled to measure the carbon (C), nitrogen (N) and phosphorus (P) content in their six organs (current-year leaves, perennial leaves, current-year twigs, perennial twigs, absorptive roots and transport roots).

Important findings (1) Current-year organs exhibited higher N and P contents but lower C:N, C:P, and N:P ratios compared to perennial organs. Stoichiometric traits showed considerable intraspecific variation across organs of different ages, with organ age identified as the primary factor influencing these differences. (2) Standardized major axis analysis revealed that perennial leaves had higher P content per unit mass than current-year leaves, whereas current-year twigs and absorptive roots showed higher C, N and P contents than their perennial counterparts (perennial twigs and transport roots). (3) Additionally, absorptive roots had significantly higher C, N and P contents per unit mass than both current-year leaves and twigs. (4) Redundancy analysis indicated that the main environmental drivers of stoichiometric traits in current-year and perennial leaves were mean annual temperature and soil P content, respectively. The most significant environmental factors affecting the stoichiometric traits of current-year and perennial twigs were annual precipitation and soil P content, respectively; while the main environmental factors influencing absorptive and transport roots were soil moisture and soil N content, respectively. This study reveals the resource allocation strategies and environmental adaptability of leaves, twigs, and roots across different age classes in P. tabuliformis. The findings demonstrate that the studies on the stoichiometry of evergreen species should fully consider age-related differences among organs, which is essential for elucidating their nutrient allocation strategies and environmental adaption mechanisms.

Key words: intraspecific variation, Pinus tabuliformis, leaf, twig, root, age, stoichiometric characteristic

于江珊, 许浩, 郭永忠, 侯继华. 器官年龄在油松叶、枝、根化学计量特征变异以及协调中的重要作用. 植物生态学报, 2026, 50(2): 306-317. DOI: 10.17521/cjpe.2025.0097

YU Jiang-Shan, XU Hao, GUO Yong-Zhong, HOU Ji-Hua. Important role of organ age in variation and coordination of stoichiometry in Pinus tabuliformis leaves, twigs and roots. Chinese Journal of Plant Ecology, 2026, 50(2): 306-317. DOI: 10.17521/cjpe.2025.0097

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2025.0097

https://www.plant-ecology.com/CN/Y2026/V50/I2/306

图/表 5

图1 9个天然油松林中, 油松叶、枝和根的碳(C)、氮(N)、磷(P)化学计量特征分布的小提琴图。不同小写字母表示不同器官间的显著差异(p < 0.05)。

Fig. 1 Violin plots showing the distribution of carbon (C), nitrogen (N), and phosphorus (P) stoichiometry in leaves, twigs and roots of P. tabuliformis across nine natural forests. Different lowercase letters indicate significant differences among organs (p < 0.05).

图2 油松当年生器官与多年生器官碳(C)、氮(N)、磷(P)化学计量特征的相对频率(A-C)以及利用嵌套线性混合模型对叶、枝和根进行的方差分解(D-F)。A和D为叶, B和E为枝, C和F为根。A-C中的垂直线表示每个年龄组的均值, 蓝色字体和黄色字体分别表示当年生和多年生器官的变异系数。D-F中的方差分解分为4个嵌套级别: 地点、样地、个体和器官年龄。C, 碳含量; N, 氮含量; P, 磷含量。

Fig. 2 Stoichiometric characteristics of the carbon (C), nitrogen (N) and phosphorus (P) in current-year and perennial organs of P. tabuliformis: Relative frequency (A-C) and variance partitioning using nested linear mixed models for leaves, branches, and roots (D-F). A and D represent leaves, B and E represent twigs, C and F represent roots. The vertical lines in A-C indicate the mean values for each age group, with blue and yellow font denoting the coefficient of variation for current-year and perennial organs, respectively. In D-F, the variance partitioning is divided into four nested levels: site, plot, individual, and organ age. C, carbon content; N, nitrogen content; P, phosphorus content.

图3 油松当年生器官与多年生器官碳(C)、氮(N)、磷(P)含量的标准主轴回归分析。线条仅在关系显著时(p < 0.05)显示。图下方标注了各器官的异速生长斜率及其95%置信区间。不同小写字母表示不同器官间差异显著(p < 0.05); ns, 不存在共同斜率。

Fig. 3 Standardized major axis regression analysis of carbon, nitrogen, phosphorus content between current-year and perennial organs in P. tabuliformis. Regression lines are shown only when the relationship is significant (p < 0.05). The allometric slopes with 95% confidence intervals are indicated in the lower corner of each panel. Different lowercase letters represent significant difference among organs (p < 0.05); ns, no common slope.

图4 油松叶、枝和根化学计量特征与环境因子的冗余分析(RDA)。A, 当年生叶。B, 当年生枝。C, 吸收根。D, 多年生叶。E, 多年生枝。F, 运输根。Current_C, 当年生器官碳含量; Current_N, 当年生器官氮含量; Current_P, 当年生器官磷含量; Current_C:N, 当年生器官碳氮比; Current_C:P, 当年生器官碳磷比; Current_N:P, 当年生器官氮磷比; Perennial_C, 多年生器官碳含量; Perennial_N, 多年生器官氮含量; Perennial_P, 多年生器官磷含量; Perennial_C:N, 多年生器官碳氮比; Perennial_C:P, 多年生器官碳磷比; Perennial_N:P, 多年生器官氮磷比; MAP, 年降水量; MAT, 年平均气温; SBD, 土壤密度; SMC, 土壤含水率; SNC, 土壤全氮含量; SPC, 土壤全磷含量。

Fig. 4 Redundancy analysis (RDA) of stoichiometry in leaves, twigs and roots of P. tabuliformis with environmental factors. A, Current-year leaves. B, Current-year twigs. C, absorptive roots. D, Perennial leaves. E, Perennial twigs. F, transport roots. Current_C, current-year organ carbon content; Current_N, current-year organ nitrogen content; Current_P, current-year organ phosphorus content; Current_C:N, carbon-to-nitrogen ratio of current-year organ; Current_C:P, carbon-to-phosphorus ratio of current-year organ; Current_N:P, nitrogen-to-phosphorus ratio of current-year organ; Perennial_C, perennial organ carbon content; Perennial_N, perennial organ nitrogen content; Perennial_P, perennial organ phosphorus content; Perennial_C:N, carbon-nitrogen ratio of perennial organ; Perennial_C:P, carbon-phosphorus ratio of perennial organ; Perennial_N:P, nitrogen-phosphorus ratio of perennial organ. MAP, mean annual precipitation; MAT, mean annual air temperature; SBD, soil bulk density; SMC, soil moisture content; SNC, soil total nitrogen content; SPC, soil total phosphorus content.

表1 环境因子对油松叶、枝和根化学计量特征的解释率

Table 1 Proportion of variance explained by environmental factors for the stoichiometry of leaf, twig and root of Pinus tabuliformis

年龄组 Age group	叶 Leaf			枝 Twig			根 Root
年龄组 Age group	环境因子 Environmental factor	解释率 Contribution rate (%)	p	环境因子 Environmental factor	解释率 Contribution rate (%)	p	环境因子 Environmental factor	解释率 Contribution rate (%)	p
当年生 Current-year	SBD	0.25	0.91	SBD	4.52	0.17	SBD	6.94	0.07
	SNC	4.24	0.18	SNC	4.80	0.13	SNC	2.15	0.40
	SPC	2.66	0.34	SPC	14.60	0.002	SPC	2.15	0.43
	SMC	3.78	0.24	SMC	5.81	0.09	SMC	32.59	<0.01
	MAT	10.50	<0.05	MAT	17.20	<0.01	MAT	7.72	0.04
	MAP	1.90	0.47	MAP	18.67	<0.01	MAP	0.61	0.79
多年生 Perennial	SBD	5.23	0.11	SBD	0.02	0.99	SBD	16.46	<0.01
	SNC	34.03	<0.01	SNC	13.17	<0.01	SNC	34.20	<0.01
	SPC	47.41	<0.01	SPC	37.12	<0.01	SPC	24.41	<0.01
	SMC	4.07	0.20	SMC	10.72	<0.01	SMC	28.47	<0.01
	MAT	5.01	0.14	MAT	16.48	<0.01	MAT	32.99	<0.01
	MAP	11.53	<0.01	MAP	12.03	<0.01	MAP	4.52	0.16

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器官年龄在油松叶、枝、根化学计量特征变异以及协调中的重要作用

Important role of organ age in variation and coordination of stoichiometry in Pinus tabuliformis leaves, twigs and roots

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