前期氮添加对无梗花栎幼苗干旱响应中地上-地下碳氮分配动态的影响

doi:10.17521/cjpe.2024.0218

植物生态学报 ›› 2025, Vol. 49 ›› Issue (9): 1527-1542.DOI: 10.17521/cjpe.2024.0218 cstr: 32100.14.cjpe.2024.0218

前期氮添加对无梗花栎幼苗干旱响应中地上-地下碳氮分配动态的影响

冯梅¹, 欧阳胜男¹^,²^,^*(), 李迈和²^,³, 周晓倩¹, 铁烈华¹, 申卫军⁴, 段洪浪¹

¹贵州大学贵州省森林资源与环境研究中心/贵州省高原山地林木培育重点实验室/贵州大学林学院, 贵阳 550025
²Swiss Federal Institute for Forest, Snow and Landscape Research, Birmensdorf 8903, Switzerland
³河北大学生命科学学院, 河北保定 071000
⁴广西大学林学院, 南宁 530004
⁵Institute of Terrestrial Ecosystems, Swiss Federal Institute of Technology, Zurich 8902, Switzerland

收稿日期:2024-07-04 接受日期:2025-01-20 出版日期:2025-09-20 发布日期:2025-04-14
通讯作者: ^*欧阳胜男 (snouyang@gzu.edu.cn)
基金资助:
国家自然科学基金(32360259);国家自然科学基金(32460379);贵州省科技计划(黔科合基础-ZK[2023]一般110);贵州省科技计划(ZK[2022]一般101)

Effects of previous nitrogen addition on aboveground and belowground carbon and nitrogen allocation dynamics in drought-exposed sessile oak seedlings

FENG Mei¹, OUYANG Sheng-Nan¹^,²^,^*(), Matthias SAURER², LI Mai-He²^,³, ZHOU Xiao-Qian¹, TIE Lie-Hua¹, SHEN Wei-Jun⁴, DUAN Hong-Lang¹, Arthur GESSLER²^,⁵

¹Institute for Forest Resources and Environment of Guizhou, Guizhou Key Laboratory of Forest Cultivation in Plateau Mountain, College of Forestry，Guizhou University, Guiyang 550025, China
²Swiss Federal Institute for Forest, Snow and Landscape Research, Birmensdorf 8903, Switzerland
³School of Life Science, Hebei University, Baoding, Hebei 071000, China
⁴College of Forestry, Guangxi University, Nanning 530004, China
⁵Institute of Terrestrial Ecosystems, Swiss Federal Institute of Technology, Zurich 8902, Switzerland

Received:2024-07-04 Accepted:2025-01-20 Online:2025-09-20 Published:2025-04-14
Supported by:
National Natural Science Foundation of China(32360259);National Natural Science Foundation of China(32460379);Guizhou Provincial Science and Technology Projects(ZK[2023]YIBAN110);Guizhou Provincial Science and Technology Projects(ZK[2022]YIBAN101)

摘要/Abstract

摘要：

干旱打破树木的碳(C)平衡过程是诱发树木干旱死亡的主要因素, 这一过程可能受到土壤养分(如氮(N))有效性的调控。然而, 氮添加对树木干旱响应过程中地上-地下部分碳和氮利用、各器官碳氮耦合关系的影响仍不清楚。该研究通过为期两年的微宇宙实验, 在第一个生长季对无梗花栎(Quercus petraea)进行氮添加处理, 随后在第二个生长季进行干旱处理, 并在干旱前进行¹⁵N标记、干旱中进行¹³C标记, 双标记后经过连续3次破坏性取样研究氮添加对无梗花栎响应持续干旱过程中碳氮分配动态的影响。主要研究结果: 1)前期氮添加促进了无梗花栎地上部分光合碳和氮分配, 也促进了根系氮吸收, 改变了植株地上-地下新吸收碳和氮的关系, 降低了各器官非结构性碳水化合物(NSC)含量; 2)干旱对无梗花栎地上-地下碳氮分配过程、各器官碳和氮关系的影响较小, 但降低了各器官NSC含量; 3)干旱第40-73天过程中, 前期氮添加下的无梗花栎表现出将吸收的氮和光合碳逐渐向地下器官转移的趋势。综上, 无梗花栎通过调控自身对碳和氮的利用策略, 具有较好的干旱适应能力, 而前期氮添加很可能会增加无梗花栎的干旱敏感性。

关键词: 氮添加, 干旱, 碳分配, 氮分配, ¹³C同位素标记, ¹⁵N同位素标记

Abstract:

Aims Drought effects on the carbon (C) balance are considered the major factor of tree mortality and are assumed to be regulated by soil nutrient (e.g., nitrogen (N)) availability. However, the effects of nitrogen addition on trees’ carbon and nitrogen distribution between aboveground and belowground and the coupling between carbon and nitrogen relations in various organs in response to drought are still unclear in trees.
Methods A two-year full factorial microcosm experiment was set up with sessile oak (Quercus petraea). Nitrogen addition was performed in the first year, and a drought treatment was conducted in the second year. Isotope ¹⁵N and ¹³C labelling were carried out before drought and during drought, respectively. Three consecutive samplings were conducted after the dual labelling with ¹³C and ¹⁵N in the second year, and the effects of nitrogen addition on carbon and nitrogen allocation dynamics during progressive drought were tested.
Important findings Our results showed that previous nitrogen addition promoted photosynthetic carbon fixation and nitrogen allocation, increased root nitrogen uptake, reduced the non-structural carbohydrates (NSC) contents in all organs and changed the relationships of carbon and nitrogen in aboveground and belowground organs. In contrast, drought had minor effects on nitrogen and carbon allocation between aboveground and belowground and the relationship of carbon with nitrogen in all organs (represented by the ratio of ¹³C to ¹⁵N in all organs). Drought only significantly reduced the content of NSC. During drought (from day 40 to 73), previous nitrogen addition led sessile oak to prioritise belowground carbon and nitrogen allocation. Our results indicate that sessile oak can change its carbon and nitrogen allocation strategies to adapt to drought, while previous nitrogen addition may increase its drought sensitivity.

Key words: nitrogen addition, drought, carbon allocation, nitrogen allocation, ¹³C isotope labelling, ¹⁵N isotope labelling

冯梅, 欧阳胜男, 李迈和, 周晓倩, 铁烈华, 申卫军, 段洪浪. 前期氮添加对无梗花栎幼苗干旱响应中地上-地下碳氮分配动态的影响. 植物生态学报, 2025, 49(9): 1527-1542. DOI: 10.17521/cjpe.2024.0218

FENG Mei, OUYANG Sheng-Nan, Matthias SAURER, LI Mai-He, ZHOU Xiao-Qian, TIE Lie-Hua, SHEN Wei-Jun, DUAN Hong-Lang, Arthur GESSLER. Effects of previous nitrogen addition on aboveground and belowground carbon and nitrogen allocation dynamics in drought-exposed sessile oak seedlings. Chinese Journal of Plant Ecology, 2025, 49(9): 1527-1542. DOI: 10.17521/cjpe.2024.0218

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2024.0218

https://www.plant-ecology.com/CN/Y2025/V49/I9/1527

图/表 10

图1 氮添加和干旱处理下无梗花栎净光合速率和叶片凌晨水势的动态变化(平均值±标准误, n = 3)。N0, 不施氮处理; N+, 氮添加处理。不同小写字母表示不同处理时间具有显著差异(p < 0.05)。

Fig. 1 Dynamics of net photosynthesis and predawn leaf water potential in sessile oak under nitrogen (N) addition and drought (mean ± SE, n = 3). N0, no N addition treatment; N+, N addition treatment. Different lowercase letters indicate significant differences among the different processing time (p < 0.05).

表1 氮添加、干旱、采样时间及其交互作用对无梗花栎各器官13C和15N含量比值、净光合速率和叶片凌晨水势的影响(p值)

Table 1 Effects of nitrogen addition, drought, sampling time, and their interaction on the ratio of 13C and 15N content in all orangs, net photosynthetic rate, and predawn leaf water potential of sessile oak (p value)

因素 Factor	¹³C:¹⁵N				净光合速率 Net photosynthetic rate (µmol·m^-2·s^-1)	叶片凌晨水势 Predawn leaf water potential (MPa)
因素 Factor	叶 Leaf	茎 Stem	细根 Fine root	粗根 Coarse root	净光合速率 Net photosynthetic rate (µmol·m^-2·s^-1)	叶片凌晨水势 Predawn leaf water potential (MPa)
N	0.009	0.008	<0.001	<0.001	0.136	0.732
D	0.993	0.850	0.199	0.853	<0.001	<0.001
Time	0.302	0.356	0.011	0.002	<0.001	0.004
D × Time	0.867	0.796	0.167	0.692	0.116	0.027
N × Time	0.310	0.273	0.139	0.015	<0.001	0.193
D × N	0.888	0.917	0.931	0.655	0.901	0.767
D × N × Time	0.854	0.719	0.210	0.773	0.240	0.049

图2 氮添加和干旱下无梗花栎各器官13C相对分配比例(平均值±标准误, n = 3)。N0, 不施氮处理; N+, 氮添加处理。不同大写字母表示不施氮和氮添加处理间具有显著差异(p < 0.05), 不同小写字母表示3个取样时间点(复水后的第15、40、73天)之间具有显著差异(p < 0.05)。

Fig. 2 Relative 13C allocation to different organs in sessile oak under nitrogen (N) addition and drought at three harvest times (mean ± SE, n = 3). N0, no N addition; N+, N addition treatment. Different uppercase letters indicate significant differences between the no N addition and the N addition treatment (p < 0.05), and different lowercase letters indicate significant differences among the three sampling time points (the 15, 40 and 73 days after rewetting) (p < 0.05).

表2 氮添加、干旱、采样时间及其交互作用对无梗花栎各器官13C、15N相对分配比例及13C、15N地上-地下比值的影响(p值)

Table 2 Effects of nitrogen addition, drought, sampling time, and their interaction on the relative allocation ratio of 15N, 13C, and the ratio of aboveground to belowground allocation in sessile oak organs (p value)

因素 Factor	叶相对分配比例 Leaf relative allocation ratio		茎相对分配比例 Stem relative allocation ratio		细根相对分配比例 Fine root relative allocation ratio
因素 Factor	¹³C	¹⁵N	¹³C	¹⁵N	¹³C	¹⁵N	¹³C	¹⁵N	¹³C	¹⁵N
N	0.948	0.124	0.025	0.159	0.027	0.001	0.154	0.736	0.003	0.039
D	0.760	0.505	0.756	0.263	0.978	0.283	0.590	0.758	0.188	0.072
Time	0.974	0.251	0.187	0.450	0.017	0.089	0.722	0.588	0.063	0.015
D × Time	0.942	0.878	0.890	0.413	0.461	0.148	0.523	0.967	0.039	0.038
N × Time	0.549	0.609	0.352	0.127	0.191	0.319	0.255	0.911	0.003	<0.001
D × N	0.082	0.020	0.838	0.127	0.657	0.764	0.321	0.618	0.151	0.432
D × N × Time	0.191	0.036	0.905	0.508	0.707	0.347	0.730	0.260	0.594	<0.001

图3 氮添加和干旱下3个取样时间点无梗花栎各器官15N相对分配比例(平均值±标准误, n = 3)。N0, 不施氮处理; N+, 氮添加处理。不同大写字母表示不施氮和氮添加处理间具有显著差异(p < 0.05), 不同小写字母表示3个取样时间点之间具有显著差异(p < 0.05)。

Fig. 3 Relative allocation ratio of 15N in organs of sessile oak under nitrogen (N) addition and drought at three harvest times (mean ± SE, n = 3). N0, no N addition; N+, N addition treatment. Different uppercase letters indicate significant differences between the no N addition and the N addition treatment (p < 0.05), and different lowercase letters indicate significant differences among the three time pointsof sampling (p < 0.05).

表3 氮添加、干旱、采样时间及其交互作用对无梗花栎各器官非结构性碳水化合物(NSC)含量和根系氮吸收速率的影响(p值)

Table 3 Effects of nitrogen addition, drought, sampling time, and their interaction on non-structural carbohydrates (NSC) content and root nitrogen uptake rate in various organs of sessile oak (p value)

因素 Factor	叶NSC含量 Leaf NSC content (%)	茎NSC含量 Stem NSC content (%)	细根NSC含量 Fine root NSC content (%)	粗根NSC含量 Coarse root NSC content (%)	根系氮吸收速率 Root nitrogen uptake rate (%)
N	0.016	0.004	0.032	0.261	<0.001
D	0.344	0.153	0.030	<0.001	0.136
Time	0.008	0.003	0.228	<0.001	<0.001
D × Time	0.563	0.250	0.022	<0.001	0.116
N × Time	0.205	0.929	0.726	0.072	<0.001
D × N	0.797	0.096	0.842	0.351	0.901
D × N × Time	0.863	0.103	0.684	0.700	0.240

图4 氮添加和干旱下3个取样时间点无梗花栎各器官非结构性碳水化合物含量(平均值±标准误, n = 3)。N0, 不施氮处理; N+, 氮添加处理。不同小写字母表示3个取样时间点之间具有显著差异(p < 0.05); *表示不施氮和氮添加处理间具有显著性差异(p < 0.05)。

Fig. 4 Tissues’ non-structural carbohydrate content in sessile oak under nitrogen (N) addition and drought at three harvest times (the 15, 40 and 73 days after rewetting) (mean ± SE, n = 3). N0, no N addition; N+, N addition treatment. Different lowercase letters indicate significant differences among the three sampling time points (p < 0.05); * indicate significant differences between the no N addition and the N addition treatment under the two watering regimes (p < 0.05).

图5 氮添加和干旱下无梗花栎3个取样时间点的根系氮吸收速率(平均值±标准误, n = 3)。DN+, 干旱条件下氮添加组; DN0, 干旱条件下不施氮组; WN+, 正常浇水条件下氮添加组; WN0, 正常浇水条件下不施氮组。不同大写字母表示不施氮和施氮间具有显著差异(p < 0.05), 不同小写字母表示3个取样时间点之间具有显著差异(p < 0.05)。

Fig. 5 Root nitrogen uptake rate in sessile oak under nitrogen (N) additions and drought at three harvest times (mean ± SE, n = 3). DN+, drought-exposed sessile oak with N addition; DN0, the drought-exposed sessile oak without N addition; WN+, the well-watered sessile oak with N addition; WN0, the well-watered sessile oak without N addition. Different uppercase letters indicate significant differences between the no N addition and the N addition treatment (p < 0.05), and different lowercase letters indicate significant differences among the three time points of sampling (p < 0.05).

图6 氮添加和干旱下无梗花栎3个取样时间点各器官的13C与15N含量比值(平均值±标准误, n = 3)。N0, 不施氮处理; N+, 氮添加处理。不同的小写字母表示3个取样时间点之间具有显著差异(p < 0.05), *表示不施氮和氮添加处理间具有显著性差异(p < 0.05)。

Fig. 6 Rate of 13C to 15N content in sessile oak under nitrogen (N) additions and drought at three harvest times (mean ± SE, n = 3). N0, no N addition; N+, N addition treatment. Different lowercase letters indicate significant differences among the three sampling time points (p < 0.05), * indicate significant differences between the no N addition and the N addition treatment under the two watering regimes (p < 0.05).

图7 前期氮添加、干旱及其两者的共同作用对无梗花栎地上-地下碳氮耦合关系的影响。表示无显著影响; 表示抑制作用; 表示促进作用, 灰色箭头表示氮从地下向地上运输, 白色箭头表示光合碳从地上向地下运输, 中间的闭环表示地上部分光合作用碳合成和分配与地下根系氮吸收和分配形成一个耦联过程。

Fig. 7 Effects of previous nitrogen addition, drought, and their combination on the coupling relationship between aboveground and underground carbon (C) and nitrogen (N) in sessile oak. represent no significant effects, inhibition and promotion, respectively. Grey arrows indicate nitrogen transport from belowground to aboveground, white arrows indicate photosynthetic carbon transport from aboveground to belowground, and the closed loop in the middle indicates that photosynthetic carbon synthesis and distribution in the aboveground part and nitrogen absorption and distribution in the belowground root system form a coupled process.

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前期氮添加对无梗花栎幼苗干旱响应中地上-地下碳氮分配动态的影响

Effects of previous nitrogen addition on aboveground and belowground carbon and nitrogen allocation dynamics in drought-exposed sessile oak seedlings

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