臭氧浓度升高和氮添加对杨树不同叶位和生长时期光合特性的影响

doi:10.17521/cjpe.2024.0352

植物生态学报 ›› 2025, Vol. 49 ›› Issue (11): 1919-1933.DOI: 10.17521/cjpe.2024.0352 cstr: 32100.14.cjpe.2024.0352

臭氧浓度升高和氮添加对杨树不同叶位和生长时期光合特性的影响

田彤彤¹, 尚博¹^,²^,^*(), 徐彦森¹^,², 袁相洋¹^,², 刘硕³, 冯兆忠¹^,²

¹南京信息工程大学生态与应用气象学院, 中国气象局生态系统碳源汇重点开放实验室, 南京 210044
²南京信息工程大学气象灾害预报预警与评估协同创新中心, 南京 210044
³浙江工业大学浙江碳中和创新研究院, 浙江省碳减排与碳监测技术国际科技合作基地, 杭州 310014

收稿日期:2024-10-10 接受日期:2025-01-10 出版日期:2025-11-20 发布日期:2025-11-20
通讯作者: *尚博(shangbo@nuist.edu.cn)
基金资助:
国家重点研发计划(2023YFC3709403);江苏省碳达峰碳中和科技创新专项(前沿基础)(BK20220020)

Effects of elevated ozone concentration and nitrogen addition on the photosynthetic characteristics of poplar at different leaf positions and growth stages

TIAN Tong-Tong¹, SHANG Bo¹^,²^,^*(), XU Yan-Sen¹^,², YUAN Xiang-Yang¹^,², LIU Shuo³, FENG Zhao-Zhong¹^,²

¹Key Laboratory of Ecosystem Carbon Source and Sink, China Meteorological Administration (ECSS-CMA), School of Applied Meteorology, Nanjing University of Information Science & Technology, Nanjing 210044, China
²Collaborative Innovation Center on Forecast and Evaluation of Meteorological Disasters(CIC-FEMD), Nanjing University of Information Science & Technology, Nanjing 210044, China
³Zhejiang Carbon Neutral Innovation Institute, Zhejiang International Cooperation Base for Science and Technology on Carbon Emission Reduction and Monitoring, Zhejiang University of Technology, Hangzhou 310014, China

Received:2024-10-10 Accepted:2025-01-10 Online:2025-11-20 Published:2025-11-20
Supported by:
National Key R&D Program of China(2023YFC3709403);Special Technology Innovation Fund of Carbon Peak and Carbon Neutrality in Jiangsu Province(BK20220020)

摘要/Abstract

摘要： 臭氧(O₃)污染和氮添加对植物光合作用的复合影响仍存在争议, 这可能与测定时期、时间及叶片位置等因素有关。该研究利用开顶式气室, 以杂交杨树(Populus euramericana cv. ‘74/76’)扦插苗为研究对象, 设置了两个O₃浓度处理(NF: 环境大气; NF45: 环境大气+ 45 nmol·mol^-1 O₃), 并在每个O₃处理下嵌套设置4个氮添加处理, 旨在探究杨树整个生长季中不同的时期、一天内不同时间及不同的叶片位置下光合参数对O₃和氮添加的响应差异。结果表明, 在4个氮处理下, 与NF处理相比, NF45处理显著地降低了整个生长季光合速率(P_n)、气孔导度(g_s)和叶绿素含量(SPAD), 分别降低了44.2%、18.2%和24.7%, 并且增加了胞间CO₂浓度(C_i) 9.0%, 表明O₃对杨树叶片P_n的降低主要是由非气孔因素限制的。比较生长季不同时期的结果发现, O₃处理对杨树生长中后期P_n和SPAD的抑制作用显著强于前期。同时, 不同叶位光合参数对O₃的响应也存在差异, NF45处理显著降低了下部叶片的P_n和SPAD, 而对最上部叶位的P_n和SPAD则有所增加, 表明植物在应对O₃胁迫时表现出一定的补偿效应。一天内不同时间的测定结果表明, O₃导致杨树叶片P_n降低的幅度不同, 但O₃和时间对杨树叶片光合参数不存在统计学上显著的交互影响。氮添加下整个生长季杨树叶片的P_n和SPAD显著升高, C_i显著降低, 但g_s没有显著的变化。氮添加对光合的促进作用在杨树的不同生长时期和不同叶位上无显著差异。此外, O₃和氮添加对所有光合参数都未观察到显著的交互效应, 表明氮添加并没有缓解O₃对杨树光合的负面影响。综上所述, 评估O₃污染对森林生态系统碳固定能力影响时, 考虑植物叶片生长阶段和位置的异质性是十分必要的, 该研究也为在空气污染背景下优化人工林氮肥管理措施提供科学依据。

关键词: 臭氧, 氮添加, 生长时期, 日变化, 叶片位置, 杨树

Abstract:

Aims The combined effects of ozone (O₃) pollution and nitrogen (N) addition on plant photosynthesis are still controversial, which may be related to factors such as measurement period, time, and leaf position. This study mainly explored the differences in the response of photosynthetic parameters of poplar to O₃ and N addition at different growth stages, diurnal variation, and leaf positions, in order to comprehensively understand the combined effects of O₃ pollution and N addition on plant carbon sequestration.

Methods The cuttings of hybrid poplar clone 107 (Populus euramericana cv. ‘74/76’) were exposed to six open top chambers. Two O₃ concentration treatments were set up (NF, non-filtered ambient air; NF45, NF + 45 nmol·mol^-1 O₃), with four N addition treatments nested under each O₃ treatment (N0, no N added; N50, N0 + 50 kg N·hm^-2·a^-1; N100, N0 + 100 kg N·hm^-2·a^-1; N200, N0 + 200 kg N·hm^-2·a^-1). The photosynthetic parameters of poplar at different growth stages, daily times, and leaf positions were measured.

Important findings When averaged across four N treatments, NF45 treatment significantly reduced the entire growth season photosynthetic rate (P_n), stomatal conductance (g_s), and chlorophyll content (SPAD) by 44.2%, 18.2%, and 24.7% compared with NF treatment, respectively. However, O₃ increased the intercellular CO₂ concentration (C_i) by 9.0%, indicating that the reduction of P_n by O₃ was mainly limited by non-stomatal factors. The results showed that NF45 treatment had a greater reduction in leaf P_n and SPAD at the middle and late stages of growth than at the early stages. The response of P_n and SPAD to O₃ varied among different leaf positions. O₃ significantly reduced P_n and SPAD at the lower leaf position, while O₃ increased P_n and SPAD at the topmost leaf position, indicating a compensatory effect of plants in response to O₃ stress. O₃ caused varying degrees of decrease in P_n of poplar at different times within a day, but there was no statistically significant interaction between O₃ and time on the photosynthetic parameters. The N addition significantly increased the leaf P_n and SPAD of poplar throughout the entire growing season, with no significant effect on g_s, but significantly reduced C_i. There was no significant difference in the promotion of photosynthesis by N addition at different growth stages and leaf positions. In addition, no significant interaction effect was observed between O₃ and N addition on all photosynthetic parameters, indicating that N addition did not alleviate the negative impact of O₃ on poplar photosynthesis.

Key words: ozone, nitrogen addition, growth period, diurnal dynamics, leaf position, poplar

田彤彤, 尚博, 徐彦森, 袁相洋, 刘硕, 冯兆忠. 臭氧浓度升高和氮添加对杨树不同叶位和生长时期光合特性的影响. 植物生态学报, 2025, 49(11): 1919-1933. DOI: 10.17521/cjpe.2024.0352

TIAN Tong-Tong, SHANG Bo, XU Yan-Sen, YUAN Xiang-Yang, LIU Shuo, FENG Zhao-Zhong. Effects of elevated ozone concentration and nitrogen addition on the photosynthetic characteristics of poplar at different leaf positions and growth stages. Chinese Journal of Plant Ecology, 2025, 49(11): 1919-1933. DOI: 10.17521/cjpe.2024.0352

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2024.0352

https://www.plant-ecology.com/CN/Y2025/V49/I11/1919

图/表 9

表1 臭氧处理、氮添加和杨树不同生长时期以及交互作用对光合参数影响的重复测量方差分析结果

Table 1 RM-ANOVA results (p values) of the interaction between O3 treatment, nitrogen addition, different growth stages and their interaction on photosynthetic parameters of poplar

变量 Parameter	净光合速率 Net photosynthetic rate	气孔导度 Stomatal conductance	胞间CO₂浓度 Intercellular CO₂concentration	叶绿素相对值 Relative chlorophyll content
臭氧 O₃	<0.01	<0.01	<0.01	<0.01
氮处理 Nitrogen (N)	<0.01	0.33	<0.01	<0.01
时期 Date	<0.01	<0.01	<0.01	<0.01
臭氧×氮处理 O₃ × N	0.49	0.72	0.37	0.65
臭氧×时期 O₃ × Date	<0.01	0.32	<0.01	<0.01
氮处理×时期 N × Date	0.40	0.10	0.18	0.02
臭氧×氮处理×时期 O₃ × N × Date	0.70	0.63	0.90	0.02

图1 不同臭氧和氮添加处理对杨树不同生长时期叶片光合速率(Pn)的影响(平均值±标准差)。NF表示环境大气, NF45表示环境大气+ 45 nmol·mol-1 O3; N0表示不添加氮, N50、N100和N200分别表示添加50、100和200 kg N·hm-2·a-1。*, p < 0.05; **, p < 0.01。

Fig. 1 Effects of different ozone and nitrogen (N) addition treatments on the leaf photosynthetic rate (Pn) of poplar at different growth stages (mean ± SD). NF, non-filtered ambient air; NF45, NF + 45 nmol·mol-1 O3. N0, no N added; N50, N0 + 50 kg N·hm-2·a-1; N100, N0 + 100 kg N·hm-2·a-1; N200, N0 + 200 kg N·hm-2·a-1. *, p < 0.05; **, p < 0.01.

图2 不同臭氧和氮添加处理对杨树不同生长时期叶片气孔导度(gs)的影响(平均值±标准差)。NF表示环境大气, NF45表示环境大气+ 45 nmol·mol-1 O3; N0表示不添加氮, N50、N100和N200分别表示添加50、100和200 kg N·hm-2·a-1。*, p < 0.05; **, p < 0.01。

Fig. 2 Effects of different ozone and nitrogen (N) addition treatments on the stomatal conductance (gs) of poplar at different growth stages (mean ± SD). NF, non-filtered ambient air; NF45, NF + 45 nmol·mol-1 O3. N0, no N added; N50, N0 + 50 kg N·hm-2·a-1; N100, N0 + 100 kg N·hm-2·a-1; N200, N0 + 200 kg N·hm-2·a-1. *, p < 0.05; **, p < 0.01.

图3 不同臭氧和氮添加处理对杨树不同生长时期叶片胞间CO2浓度(Ci)的影响(平均值±标准差)。NF表示环境大气, NF45表示环境大气+ 45 nmol·mol-1 O3; N0表示不添加氮, N50、N100和N200分别表示添加50、100和200 kg N·hm-2·a-1。*, p < 0.05; **, p < 0.01。

Fig. 3 Effects of different ozone and nitrogen (N) addition treatments on the intercellular CO2 concentration (Ci) of poplar at different growth stages (mean ± SD). NF, non-filtered ambient air; NF45, NF + 45 nmol·mol-1 O3. N0, no N added; N50, N0 + 50 kg N·hm-2·a-1; N100, N0 + 100 kg N·hm-2·a-1; N200, N0 + 200 kg N·hm-2·a-1. *, p < 0.05; **, p < 0.01.

图4 不同臭氧和氮添加处理对杨树不同生长时期叶片叶绿素相对值(SPAD)的影响(平均值±标准差)。NF表示环境大气, NF45表示环境大气+ 45 nmol·mol-1 O3; N0表示不添加氮, N50、N100和N200分别表示添加50、100和200 kg N·hm-2·a-1。*, p < 0.05; **, p < 0.01。

Fig. 4 Effects of different ozone and nitrogen (N) addition treatments on the relative chlorophyll content (SPAD) of poplar at different growth stages (mean ± SD). NF, non-filtered ambient air; NF45, NF + 45 nmol·mol-1 O3. N0, no N added; N50, N0 + 50 kg N·hm-2·a-1; N100, N0 + 100 kg N·hm-2·a-1; N200, N0 + 200 kg N·hm-2·a-1. *, p < 0.05; **, p < 0.01.

图5 整个生长季不同臭氧处理下杨树叶片相对叶绿素(SPAD)与光合速率(Pn)、气孔导度(gs)、胞间CO2浓度(Ci)的线性关系。NF表示环境大气, NF45表示环境大气+ 45 nmol·mol-1 O3。ANCOVA, 协方差分析。

Fig. 5 Linear relationship between relative chlorophyll content (SPAD) and photosynthetic rate (Pn), stomatal conductance (gs), and intercellular CO2 concentration (Ci) of poplar leaves under different ozone treatments throughout the growth season. NF, non-filtered ambient air; NF45, NF + 45 nmol·mol-1 O3. ANCOVA, analysis of covariance.

图6 不同臭氧(O3)和氮(N)添加处理对杨树叶片光合速率(Pn)、气孔导度(gs)、胞间CO2浓度(Ci)日变化(T)特征的影响(平均值±标准差)。NF表示环境大气, NF45表示环境大气+ 45 nmol·mol-1 O3; N0表示不添加氮, N200表示添加200 kg N·hm-2·a-1。蓝色*表示N0处理下两个O3处理下的差异显著; 红色*表示N200处理下两个O3处理下的差异显著; *, p < 0.05; **, p < 0.01。图中仅展示了统计学显著的p值。

Fig. 6 Effects of different ozone (O3) and nitrogen (N) addition treatments on the diurnal variation (T) of photosynthetic rate (Pn), stomatal conductance (gs), and intercellular CO2 concentration (Ci) in poplar leaves (mean ± SD). NF, non-filtered ambient air; NF45, NF + 45 nmol mol-1·O3. N0, no N added; N200, N0 + 200 kg N·hm-2·a-1. The blue asterisk indicated significant differences between the two O3 treatments under N0 treatment; The red asterisk indicated a significant difference between the two O3 treatments under N200 treatment. *, p < 0.05; **, p < 0.01. Only statistically significant p-values are shown.

图7 不同臭氧(O3)和氮(N)添加处理对杨树不同叶位(P)光合速率(Pn)的影响(平均值±标准差)。NF表示环境大气, NF45表示环境大气+ 45 nmol·mol-1 O3; N0表示不添加氮, N50、N100和N200分别表示添加50、100和200 kg N·hm-2·a-1。*, p < 0.05; **, p < 0.01。I-V分别表示从上到下的1-5、6-10、11-15、16-20、21至底部的叶片。

Fig. 7 Effects of different ozone (O3) and nitrogen (N) addition treatments on the photosynthetic rate (Pn) at different leaf positions (P) of poplar (mean ± SD). NF, non-filtered ambient air; NF45, NF + 45 nmol·mol-1 O3. N0, no N added; N50, N0 + 50 kg N·hm-2·a-1; N100, N0 + 100 kg N·hm-2·a-1; N200, N0 + 200 kg N·hm-2·a-1. *, p < 0.05; **, p < 0.01. I-V is 1-5, 6-10, 11-15, 16-20, 21 to botton leaves form top to bottom, respectively.

图8 不同臭氧(O3)和氮(N)添加处理对杨树不同叶位(P)叶绿素相对含量(SPAD)的影响(平均值±标准差)。NF表示环境大气, NF45表示环境大气+ 45 nmol·mol-1 O3; N0表示不添加氮, N50、N100和N200分别表示添加50、100和200 kg N·hm-2·a-1。*, p < 0.05; **, p < 0.01。I-V分别表示从上到下的1-5、6-10、11-15、16-20、21至底部的叶片。

Fig. 8 Effects of different ozone (O3) and nitrogen (N) addition treatments on the relative chlorophyll content (SPAD) at different leaf positions (P) of poplar (mean ± SD). NF, non-filtered ambient air; NF45, NF + 45 nmol·mol-1 O3. N0, no N added; N50, N0 + 50 kg N·hm-2·a-1; N100, N0 + 100 kg N·hm-2·a-1; N200, N0 + 200 kg N·hm-2·a-1. *, p < 0.05; **, p < 0.01. I-V is 1-5, 6-10, 11-15, 16-20, 21 to botton leaves form top to bottom, respectively.

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臭氧浓度升高和氮添加对杨树不同叶位和生长时期光合特性的影响

Effects of elevated ozone concentration and nitrogen addition on the photosynthetic characteristics of poplar at different leaf positions and growth stages

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