氮磷添加对树木生长和森林生产力影响的研究进展

doi:10.17521/cjpe.2019.0176

摘要/Abstract

摘要：

人为活动所导致的氮、磷输入和大气氮、磷沉降使生态系统中的氮、磷可利用性大幅提高, 对陆地生态系统的碳循环过程产生了显著影响。树木生长和森林生产力在全球碳循环中发挥着重要作用, 它决定着陆地碳固存的大小和方向。目前, 在全球范围内开展了很多氮、磷添加调控树木生长和森林生产力的野外控制实验, 但是研究结果并不一致, 受到多种生物、环境和实验处理条件等因素的影响。该文从野外氮添加和磷添加实验的文献数量、实验数量及其全球空间分布三个方面概述了氮、磷添加对树木生长和森林生产力影响的研究现状, 并总结了氮、磷添加实验中树木生长和森林生产力的评估方法, 包括相对生长速率和绝对增长量。基于相关的研究结果, 阐述了氮、磷添加影响树木生长和森林生产力的调控因素及其潜在影响机制, 包括气候、树木径级与林龄、植物功能性状(共生菌根类型、树木固氮属性和保守性与获得性性状)、植物和微生物相互作用关系、区域养分沉降速率和实验处理条件等。最后, 基于当前的研究进行了系统总结, 并指出今后需要加强的几个方面的研究, 以期为后续研究提供参考: 树木生长响应氮、磷添加的生理学机制, 树木各部分生长对氮、磷添加响应的权衡与分配, 植物功能性状在调节与预测树木生长响应氮、磷添加中的作用, 树木之间的竞争关系如何调控氮、磷添加对树木生长的影响, 以及开展长期的和联网的氮、磷添加对树木生长和森林生产力影响的野外控制实验。

关键词: 养分限制, 树木生长, 生态系统生产力, 植物功能性状, 植物与微生物相互作用, 氮沉降, 磷沉降

Abstract:

Nitrogen (N) and phosphorus (P) inputs induced by anthropogenic activities and atmospheric N and P deposition have largely increased the availability of soil N and P in terrestrial ecosystems, which have considerably affected terrestrial carbon cycling processes. Tree growth and productivity in forest ecosystems play an important role in global carbon cycling, and determine the magnitude and direction of terrestrial carbon sequestration. Currently, a large number of field manipulation experiments have been conducted to investigate the effects of N and/or P addition on tree growth and forest productivity, but the results from these studies were inconsistent. Such inconsistent results might be affected by multiple factors, including biological, environmental and experimental variables. Here, we reviewed the present research status of the effects of N and P addition on tree growth and forest productivity in forest ecosystems based on three aspects, including the number of publications and experiments with field N and P addition, and the global distributions of these experiments. Then, we summarized the methods for assessing tree growth and forest productivity at ecosystem level in forest ecosystems, including relative growth rate and absolute increment. According to the related results, we reviewed the regulating factors that affect tree growth and productivity, and the potential mechanisms for such factors, including climate, tree size and stand age, plant functional traits (including type of tree-associated mycorrhizal fungi, N-fixation property of trees, and conservative and acquisitive functional traits), plant-microbe interaction, ambient nutrient (i.e., N and P) deposition rate, and experimental variables. Finally, we summarized the current studies, and pointed out five aspects that are urgently needed to provide further insights in future studies, including the physiological mechanism of how tree growth responds to N and P addition, the tradeoff and allocation among growth of various parts of tree under N and P addition, the role of plant functional traits in regulating and predicting the responses of tree growth to N and P addition, how the competition among trees regulates the responses of tree growth to N and P addition, and conducting long-term and coordinated distributed field experiments investigating the effects of N and P addition on tree growth and forest productivity at the global scale.

Key words: nutrient limitation, tree growth, ecosystem productivity, plant functional traits, plant-microbe interaction, nitrogen deposition, phosphorus deposition

冯继广, 朱彪. 氮磷添加对树木生长和森林生产力影响的研究进展. 植物生态学报, 2020, 44(6): 583-597. DOI: 10.17521/cjpe.2019.0176

FENG Ji-Guang, ZHU Biao. A review on the effects of nitrogen and phosphorus addition on tree growth and productivity in forest ecosystems. Chinese Journal of Plant Ecology, 2020, 44(6): 583-597. DOI: 10.17521/cjpe.2019.0176

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2019.0176

https://www.plant-ecology.com/CN/Y2020/V44/I6/583

图/表 5

图1 氮(N)和磷(P)添加对树木生长和森林生产力影响的论文数量和实验数量。 A, 1970-2019年每年的论文数量及累积论文数量。B, 不同N、P添加处理在不同气候区森林的实验数量。N表示该研究中只有N添加; P表示该研究中只有P添加; N, P表示该研究中有单独的N和单独的P添加; N, P, NP表明该研究中包含单独的N、单独的P和N、P共同添加。

Fig. 1 Number of papers and experiments investigating the effects of nitrogen (N) and/or phosphorus (P) addition on tree growth and productivity in forest ecosystems. A, The number of papers in each year and the accumulated number of papers between the year 1970 and 2019. B, The number of experiments with different treatment of nitrogen and phosphorus addition. N represents only N addition in the case study; P represents only P addition in the case study; N, P represent only N and only P addition in the case study; N, P, NP represents only N, only P and N plus P addition in the case study.

图2 氮(N)和磷(P)添加对森林树木生长及生产力影响的案例研究全球分布图。图中的横线表示基于纬度划分的不同气候区的森林分布界限, 包括热带森林(23.5° S-23.5° N)、北方森林(46°-66° N)和温带森林(介于热带和北方森林的纬度之间)。 N表示该研究中只有N添加; P表示该研究中只有P添加; N, P表示该研究中有单独的N和单独的P添加; N, P, NP表明该研究中包含单独的N、单独的P和N、P共同添加。该图使用ArcGIS 10.2.2绘制。

Fig. 2 Global distribution of case studies investigating the effects of nitrogen (N) and/or phosphorus (P) addition on tree growth and productivity in forest ecosystems. The horizontal lines in the figure indicate the distribution boundaries of forests in different climatic regions based on latitude, including tropical forest (23.5° S-23.5° N), boreal forest (46°-66° N), and temperate forest (between the tropical and boreal latitudes). N represents only N addition in the case study; P represents only P addition in the case study; N, P represent only N and only P addition in the case study; N, P, NP represents only N, only P and N plus P addition in the case study. This map was drawn using the software ArcGIS 10.2.2.

表1 氮和磷添加对树木生长和生产力影响的评估方法

Table 1 Methods for assessing the effects of nitrogen and/or phosphorus addition on plant growth and productivity

衡量方法 Method	计算公式 Calculation equation	单位 Unit	参考文献 Reference
相对生长速率 Relative growth rate (RGR)	$RGR=\frac{\text{ln}DB{{H}_{{{t}_{2}}}}-\text{ln}DB{{H}_{{{t}_{1}}}}}{{{t}_{2}}-{{t}_{1}}}$	mm·mm^-1·a^-1	Carlson et al., 2008; Alvarez-Clare et al., 2013; Li et al., 2018
	$RGR=\frac{\text{ln}B{{A}_{{{t}_{2}}}}-\text{ln}B{{A}_{{{t}_{1}}}}}{{{t}_{2}}-{{t}_{1}}}$	mm²·mm^-2·a^-1	Tian et al., 2017
	$RGR=\frac{\text{ln}B{{P}_{{{t}_{2}}}}-\text{ln}B{{P}_{{{t}_{1}}}}}{{{t}_{2}}-{{t}_{1}}}$	kg·kg^-1·a^-1	Markewitz et al., 2012
	$RGR=\frac{B{{A}_{{{t}_{2}}}}-B{{A}_{{{t}_{1}}}}\text{ }\!\!~\!\!\text{ }}{\text{B}{{\text{A}}_{{{t}_{1}}}}}$	%	Jiang et al., 2018
绝对增长量 Absolute increment (Δ)	$\Delta =DB{{H}_{{{t}_{2}}}}-DB{{H}_{{{t}_{1}}}}$	mm·a^-1	Noguchi et al., 2013
	$\text{ }\!\!\Delta\!\!\text{ }\!\!~\!\!\text{ }=B{{A}_{{{t}_{2}}}}-B{{A}_{{{t}_{1}}}}$	m²·a^-1	Saarsalmi et al., 2012; Tian et al., 2017
	$\text{ }\!\!\Delta\!\!\text{ }\!\!~\!\!\text{ }=B{{P}_{{{t}_{2}}}}-B{{P}_{{{t}_{1}}}}$	kg·hm^-2·a^-1	Markewitz et al., 2012; Du & Fang, 2014

表2 基于胸径的树木等级划分

Table 2 Methods for assessing the effects of nitrogen and/or phosphorus addition on plant growth and productivity

胸径等级 Classes for diameter at breast height (cm)			参考文献 Reference
小树 Small tree	中树 Medium tree	大树 Large tree	参考文献 Reference
3-10	10-20	>20	Fisher et al., 2013; Jiang et al., 2018; Zou et al., 2019
5-10	10-30	>30	Alvarez-Clare et al., 2013; Tian et al., 2017
5-15		>15	Li et al., 2018
10-25		>25	Wright et al., 2018

图3 氮和磷添加对树木生长及森林生产力影响研究的总结图。 AM, 丛枝菌根; ECM, 外生菌根。

Fig. 3 Summary diagram for studying the effects of nitrogen and/or phosphorus addition on tree growth and productivity in forest ecosystems. AM, arbuscular mycorrhizal; ECM, ectomycorrhizal.

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