长期氮添加对温带落叶阔叶林优势植物叶片次生代谢产物的影响

doi:10.17521/cjpe.2024.0262

植物生态学报 ›› 2024, Vol. 48 ›› Issue (12): 1576-1588.DOI: 10.17521/cjpe.2024.0262 cstr: 32100.14.cjpe.2024.0262

长期氮添加对温带落叶阔叶林优势植物叶片次生代谢产物的影响

赵常提¹(), 夏青霖¹, 田地¹^,^*()(), 陈冰瑞¹, 朱瑞德¹, 刘宵含¹, 俞果², 吉成均²

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北京大学城市与环境学院, 生态研究中心, 地表过程分析与模拟教育部重点实验室, 北京 100871

收稿日期:2024-08-06 接受日期:2024-12-12 出版日期:2024-12-20 发布日期:2024-12-20
通讯作者: *田地(tiandi@bjfu.edu.cn)
基金资助:
北京市自然科学基金(5222018);国家自然科学基金(32271680);国家自然科学基金(31800397)

Effects of long-term nitrogen addition on leaf secondary metabolites of the dominant plant species in a temperate deciduous broad-leaved forest

ZHAO Chang-Ti¹(), XIA Qing-Lin¹, TIAN Di¹^,^*()(), CHEN Bing-Rui¹, ZHU Rui-De¹, LIU Xiao-Han¹, YU Guo², JI Cheng-Jun²

¹State Key Laboratory of Efficient Production of Forest Resources, Beijing Forestry University, Beijing 100083, China
²Institute of Ecology, College of Urban and Environmental Sciences, Key Laboratory for Earth Surface Processes of the Ministry of Education, Peking University, Beijing 100871, China

Received:2024-08-06 Accepted:2024-12-12 Online:2024-12-20 Published:2024-12-20
Contact: *TIAN Di(tiandi@bjfu.edu.cn)
Supported by:
Supported by the Beijing Municipal Natural Science Foundation(5222018);and the National Natural Science Foundation of China(32271680);and the National Natural Science Foundation of China(31800397)

摘要/Abstract

摘要： 植物次生代谢产物(PSM)与植物的生长发育、抵御生物与非生物胁迫的能力密切相关, 在植物对环境的适应策略中发挥至关重要的作用。然而, 目前森林生态系统中植物叶片PSM如何响应长期氮添加尚不清晰, 限制了对氮沉降背景下树木抗性及森林生态系统稳定性变化的理解。该研究基于连续12年的氮添加实验平台, 选取了北京东灵山温带落叶阔叶林中的乔灌层优势植物, 包括白桦(Betula platyphylla)林的优势植物白桦和毛榛(Corylus mandshurica), 以及蒙古栎(Quercus mongolica)林的优势植物蒙古栎和大花溲疏(Deutzia grandiflora), 旨在探究两种林型的土壤理化性质、4种植物的叶片酚类PSM (总酚、黄酮和单宁)含量及乔木生长对长期氮添加的响应。结果表明, 白桦林土壤水分和养分含量均高于蒙古栎林; 氮添加显著降低了白桦林的土壤pH和蒙古栎林的土壤含水量, 并增加了蒙古栎林的土壤全磷含量。整体上, 白桦林和蒙古栎林的优势植物的叶片酚类PSM含量对氮添加的响应并不一致, 表现为白桦林呈上升趋势, 而蒙古栎林则呈下降趋势, 尤其是在高氮添加处理下较为显著。4种植物的叶片酚类PSM含量与叶片养分含量存在权衡关系, 并受到土壤水分和养分的调控。此外, 氮添加后, 白桦的相对生长速率呈下降趋势, 而蒙古栎则呈上升趋势。这些结果暗示氮沉降背景下不同林型差异化的养分分配及生长-抗性平衡策略。相较于白桦林, 水分和养分相对贫瘠的蒙古栎林在长期氮沉降的影响下很可能较早出现脆弱性相关(如昆虫植食性水平增加, 病原菌丰度上升等)的问题。

关键词: 氮沉降, 温带落叶阔叶林, 植物次生代谢产物, 土壤理化性质, 养分分配策略

Abstract:

Aims Plant secondary metabolites (PSMs) are closely related to plant growth, development, and the ability to resist biotic and abiotic stresses, playing a crucial role in plant adaptation strategies to the environment. However, the response of leaf PSMs to long-term nitrogen (N) addition in forest ecosystems remains insufficiently studied, which largely limits our ability to grasp the changes in tree resistance and forest ecosystem stability under N deposition.

Methods Our study is based on an 12-year N addition experiment platform, focusing on four dominant plant species in the temperate deciduous forest of Donglingshan: including the plants Betula platyphylla and Corylus mandshurica in the birch forest and Quercus mongolica and Deutzia grandiflora in the oak forest. We investigated responses of soil physicochemical properties in two forest types, leaf phenolic PSMs (i.e., total phenolic, flavonoid, and tannin) contents of four plant species, and tree growth to long-term N addition.

Important findings The results showed that soil water and nutrient contents of birch forest were higher than those of oak forest. N addition significantly decreased soil pH in birch forest and soil water content in oak forest, and increased soil total phosphorus content in oak forest. Overall, the responses of dominant plant species to N addition exhibited contrasting trends, with leaf PSMs showing an increasing trend in the birch forest and a decreasing trend in the oak forest, particularly under high N addition treatments. The leaf phenolic PSMs contents of the four plant species exhibited a trade-off with leaf nutrient contents which was regulated by soil water and nutrient availability. Additionally, after N addition, B. platyphylla showed a decreasing trend in relative growth rate, whereas Q. mongolica showed an increasing trend. These results suggested different nutrient allocation and growth-defense balance strategies among different forest types under N deposition. Compared to the birch forest, the oak forest, which was relatively poor in water and nutrients, was likely to experience vulnerability-related issues (e.g. increased levels of insect herbivory and higher abundance of pathogens) earlier under the influence of long-term high N deposition.

Key words: nitrogen deposition, temperate deciduous broad-leaved forest, plant secondary metabolites, soil physicochemical property, nutrient allocation strategy

赵常提, 夏青霖, 田地, 陈冰瑞, 朱瑞德, 刘宵含, 俞果, 吉成均. 长期氮添加对温带落叶阔叶林优势植物叶片次生代谢产物的影响. 植物生态学报, 2024, 48(12): 1576-1588. DOI: 10.17521/cjpe.2024.0262

ZHAO Chang-Ti, XIA Qing-Lin, TIAN Di, CHEN Bing-Rui, ZHU Rui-De, LIU Xiao-Han, YU Guo, JI Cheng-Jun. Effects of long-term nitrogen addition on leaf secondary metabolites of the dominant plant species in a temperate deciduous broad-leaved forest. Chinese Journal of Plant Ecology, 2024, 48(12): 1576-1588. DOI: 10.17521/cjpe.2024.0262

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https://www.plant-ecology.com/CN/Y2024/V48/I12/1576

图/表 5

图1 氮添加对白桦林和蒙古栎林土壤理化性质的影响(n = 9个体× 3处理× 2林型= 54)。“×”表示该组数据的平均值点。每个子图右上角标注了不同林型间的统计差异p值(ns, p > 0.05; **, p < 0.01; ***, p < 0.001)。不同小写字母表示处理组间差异显著(p < 0.05)。CK, 对照, 0 kg N·hm-2·a-1; N50, 低氮, 50 kg N·hm-2·a-1; N100, 高氮, 100 kg N·hm-2·a-1。

Fig. 1 Effect of nitrogen (N) addition on soil physicochemical properties in Betula platyphylla forest and Quercus mongolica forest (n = 9 individuals × 3 treatments × 2 forests = 54). The “×” symbol represents the mean value for each group. Each subplot displays the p-values for statistical differences between the forest types in the top right corner (ns, p > 0.05; **, p < 0.01; ***, p < 0.001). Different lowercase letters indicate significant differences between treatments (p < 0.05). CK, control, 0 kg N·hm-2·a-1; N50, low nitrogen, 50 kg N·hm-2·a-1; N100, high nitrogen, 100 kg N·hm-2·a-1.

图2 氮添加对温带落叶阔叶林4种优势植物叶片酚类次生代谢产物含量的影响(n = 9个体× 3处理× 4物种= 108)。“×”表示该组数据的平均值点。不同小写字母表示处理组间差异显著(p < 0.05)。CK, 对照, 0 kg N·hm-2·a-1; N50, 低氮, 50 kg N·hm-2·a-1; N100, 高氮, 100 kg N·hm-2·a-1。

Fig. 2 Effects of nitrogen addition on leaf phenolic secondary metabolites content of four dominant plants in temperate deciduous broad-leaved forest (n = 9 individuals × 3 treatments × 4 species = 108). The “×” symbol represents the mean value for each group. Different lowercase letters indicate significant differences between different treatments (p < 0.05). CK, control, 0 kg N·hm-2·a-1; N50, low nitrogen, 50 kg N·hm-2·a-1; N100, high nitrogen, 100 kg N·hm-2·a-1.

图3 4种温带落叶阔叶优势植物的叶片酚类次生代谢产物与养分含量的主成分(PC)分析图(A)以及第一主成分(PC1)与土壤理化性质的相关性(B-F) (n = 9个体× 3处理× 4物种= 108)。不同颜色代表不同的物种。深色点代表每个物种的平均值, 浅色点代表样本点。当p < 0.05时回归线用实线表示, 否则用虚线表示。FLA, 黄酮含量; LC, 叶碳含量; LN, 叶氮含量; LP, 叶磷含量; pH, 土壤pH; STC, 土壤全碳含量; STN, 土壤全氮含量; STP, 土壤全磷含量; SWC, 土壤含水量; TA, 单宁含量; TP, 总酚含量。

Fig. 3 Principal component (PC) analysis diagram of leaf phenolic secondary metabolites and nutrient content for four dominant plants in temperate deciduous broad-leaved forest (A) and relationship between the first principal component (PC1) and soil physicochemical properties (B-F) (n = 9 individuals × 3 treatments × 4 species = 108). Different colors represent different species groups. Deep-colored dots represent the mean values of each species, while light-colored dots represent the sample points. The regression line is depicted with a solid line if p < 0.05, otherwise with a dashed line. FLA, flavonoid content; LC, leaf carbon content; LN, leaf nitrogen content; LP, leaf phosphorus content; pH, soil pH; STC, soil organic carbon content; STN, soil total nitrogen content; STP, soil total phosphorus content; SWC, soil water content; TA, tannin content; TP, total phenolic content.

图4 白桦林(A)和蒙古栎林(B)优势植物叶片酚类次生代谢产物和养分含量的主成分分析图(n = 9个体× 3处理× 2物种= 54)。不同颜色代表不同的氮添加处理。柱状图表示第一(PC1)和第二主成分(PC2)得分在不同氮添加处理间的差异, 不同小写字母表示不同处理间差异显著(p < 0.05)。FLA, 黄酮含量; LC, 叶碳含量; LN, 叶氮含量; LP, 叶磷含量; TA, 单宁含量; TP, 总酚含量。CK, 对照, 0 kg N·hm-2·a-1; N50, 低氮, 50 kg N·hm-2·a-1; N100, 高氮, 100 kg N·hm-2·a-1。

Fig. 4 Principal component analysis diagram of leaf secondary metabolites and nutrient content for Betula platyphylla forest (A) and Quercus mongolica forest (B) (n = 9 individuals × 3 treatment × 2 species = 54). Different colors represent different nitrogen treatments. Barplots show PC1 and PC2 score differences across nitrogen treatments, with significant differences indicated by different lowercase letters (p < 0.05). FLA, flavonoid content; LC, leaf carbon content; LN, leaf nitrogen content; LP, leaf phosphorus content; TA, tannin content; TP, total phenolic content. CK, control, 0 kg N·hm-2·a-1; N50, low nitrogen, 50 kg N·hm-2·a-1; N100, high nitrogen, 100 kg N·hm-2·a-1.

图5 氮添加对白桦和蒙古栎相对生长速率的影响(n = 9个体× 3处理= 27)。“×”表示该组数据的平均值点。“ns”代表每个树种的相对生长速率在不同氮添加处理间无显著差异(p < 0.05)。CK, 对照, 0 kg N·hm-2·a-1; N50, 低氮, 50 kg N·hm-2·a-1; N100, 高氮, 100 kg N·hm-2·a-1。

Fig. 5 Effects of nitrogen addition to the relative growth rate of Betula platyphylla and Quercus mongolica (n = 9 individuals × 3 treatments = 27). The “×” symbol represents the mean value for each group. “ns” indicates relative growth rate of each tree species have no significant differences among different nitrogen treatments (p < 0.05). CK, control, 0 kg N·hm-2·a-1; N50, low nitrogen, 50 kg N·hm-2·a-1; N100, high nitrogen, 100 kg N·hm-2·a-1.

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长期氮添加对温带落叶阔叶林优势植物叶片次生代谢产物的影响

Effects of long-term nitrogen addition on leaf secondary metabolites of the dominant plant species in a temperate deciduous broad-leaved forest

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