长期氮沉降对杉木人工林叶、枝氮磷养分再吸收的影响

doi:10.17521/cjpe.2018.0167

植物生态学报 ›› 2018, Vol. 42 ›› Issue (9): 926-937.DOI: 10.17521/cjpe.2018.0167

所属专题：全球变化与生态系统；生态化学计量

长期氮沉降对杉木人工林叶、枝氮磷养分再吸收的影响

沈芳芳^1,²,李燕燕^1,²,刘文飞²,段洪浪²,樊后保^2,^*(),胡良²,孟庆银³

1江西农业大学林学院, 江西省森林培育重点实验室, 南昌 330045
2南昌工程学院生态与环境科学研究所, 江西省退化生态系统修复与流域生态水文重点实验室, 南昌 330099
3沙县官庄国有林场, 福建沙县 365503

收稿日期:2018-07-16 修回日期:2018-09-10 出版日期:2018-09-20 发布日期:2019-01-15
通讯作者: 樊后保
基金资助:
国家自然科学基金(31360175);国家自然科学基金(31570444);赣鄱英才555工程和江西省主要学科学术和技术带头人项目(20162BCB22021)

Responses of nitrogen and phosphorus resorption from leaves and branches to long-term nitrogen deposition in a Chinese fir plantation

SHEN Fang-Fang^1,²,LI Yan-Yan^1,²,LIU Wen-Fei²,DUAN Hong-Lang²,FAN Hou-Bao^2,^*(),HU Liang²,MENG Qing-Yin³

1School of Forestry, Jiangxi Agricultural University, Jiangxi Key Laboratory of Silviculture, Nanchang 330045, China
2Institute of Ecology and Environmental Science, Nanchang Institute of Technology, Jiangxi Key Laboratory for Restoration of Degraded Ecosystems and Watershed Ecohydrology, Nanchang 330099, China
3Guanzhuang National Forest Farm of Shaxian, Sha Xian, Fujian 365503, China

Received:2018-07-16 Revised:2018-09-10 Online:2018-09-20 Published:2019-01-15
Contact: Hou-Bao FAN
Supported by:
Supported by the National Natural Science Foundation of China.(31360175);Supported by the National Natural Science Foundation of China.(31570444);the Gan-Po 555 Talent Project, and the Jiangxi Provincial Fund for Academic and Technical Leadership of Major Disciplines.(20162BCB22021)

摘要/Abstract

摘要：

为了解森林养分内循环对全球变化的响应, 基于长期模拟氮沉降试验, 研究了杉木(Cunninghamia lanceolata)人工林不同龄级(一年生、二年生和衰老)叶和枝的氮(N)、磷(P)养分分配及其再吸收特征, 并分析了不同模拟N沉降处理时间(7年和14年)杉木叶N、P养分再吸收差异。在12年生杉木中开展模拟N沉降试验, 以尿素(CO(NH₂)₂)为N源, 设N0、N1、N2和N3 4个处理水平, 施氮量分别为0、60、120和240 kg·hm ^-2·a ^-1, 每个处理重复3次。结果表明: (1)叶和枝在衰老过程中碳(C)、N和P含量逐渐降低, 且叶的C、N和P含量比枝高; N含量大小依次为一年生叶>二年生叶>衰老叶>一年生枝>二年生枝>衰老枝, 且N3 > N2 > N1 > N0, 而C:N则呈现相反的趋势; 衰老器官的C:N、C:P、N:P比新鲜器官高; N沉降增加了不同龄级叶和枝(除二年生叶外)的N、N:P和C:P, 但降低了P和C:N。(2)叶和枝的N、P养分再吸收率(RE_N、RE_P)随龄级的增加至衰老有规律地递减, 且RE_P > RE_N; 受长期N沉降的影响, RE_N叶(28.12%) <枝(30.00%), 而RE_P则为叶(45.82%) >枝(30.42%); 杉木叶和枝N:P与RE_N:RE_P之间存在极显著的线性相关关系。(3)随N沉降处理时间的增加, 叶RE_N呈降低态势, 各处理(N1、N2和N3)分别降低了9.85%、3.17%和11.71%; 而RE_P则明显上升, 分别增加了71.98%、42.25%和9.60%。研究结果表明: 不同器官、不同龄级的养分再吸收率随氮沉降处理的水平、处理时间而所有不同; RE_N:RE_P与N:P之间存在紧密关系。

关键词: 养分再吸收率, 生态化学计量, 杉木人工林, 长期氮沉降

Abstract:

Aims Our objectives were to investigate differences in nutrient resorption between different plant organs (leaf and branch), among plants with different life spans (one-year old, two-year old and senesced), and under different duration of nitrogen (N) deposition treatments in a Chinese fir (Cunninghamia lanceolata) plantation.

Methods The long-term N deposition experiment was conducted in a 12-year-old fir plantation of subtropical China. N deposition treatment was initiated in January 2004 until now, up-going 14 years. N deposition were designed at 4 levels of 0, 60, 120, and 240 kg·hm ^-2·a ^-1, indicated as N0, N1, N2, and N3, respectively, with 3 replicates for each treatment. The solution of CO(NH₂)₂was sprayed on the forest floor each month. In the study, we measured N and phosphorus (P) concentrations and analyzed the pattern of nutrient resorption of mature and senescing leaves and branches. The different responses of needles N and P resorption after 7- and 14-year N deposition treatments were also compared.

Important findings After 14 years of N deposition, (1) during the senescing process, leaf and branch C, N, and P content gradually decreased with increasing treatment duration, with higher content in leaf than in branch. N content decreased in the order of one-year old green leaf > two-year old green leaf > senescent leaf > one-year old living branch > two-year old living branch > senescent branch, and N3 > N2 > N1 > N0, with C:N showing the opposite trend. Senescent organs had higher C:N, N:P, and C:P than mature living organs. N deposition increased N, N:P, and C:P of mature living organs (except for the two-year old green leaf), while decreased P and C:N. (2) N resorption efficiency (RE_N) and P resorption efficiency (RE_P) of leaves and branches decreased gradually with increasing life span. RE_P was typically higher in leaf and branch than RE_N. Leaf had lower RE_N (28.12%) than branch (30.00%), but higher RE_P (45.82%) than branch (30.42%). A highly significant linear correlation existed between N:P and RE_N:RE_P in leaves and branches. (3) RE_N decreased but RE_P increased with the treatment duration of N deposition. The longer experimental duration (14 years) reduced RE_N by 9.85%, 3.17%, 11.71% under N1, N2, and N3 treatments, respectively, and increased RE_P by 71.98%, 42.25%, 9.60%, respectively, than the shorter treatment duration (7 years). In summary, the responses of essential nutrients resorption efficiency for different plant organs and life span varied with the levels and duration of N deposition treatment. RE_N:RE_P in leaf and branch were mostly driven by N:P of leaf and branch. The results highlight that nutrients resorption is significantly influenced by long-term N deposition.

Key words: nutrient resorption efficiency, ecological stoichiometry, Cunninghamia lanceolata plantation, long-term nitrogen deposition

沈芳芳, 李燕燕, 刘文飞, 段洪浪, 樊后保, 胡良, 孟庆银. 长期氮沉降对杉木人工林叶、枝氮磷养分再吸收的影响. 植物生态学报, 2018, 42(9): 926-937. DOI: 10.17521/cjpe.2018.0167

SHEN Fang-Fang, LI Yan-Yan, LIU Wen-Fei, DUAN Hong-Lang, FAN Hou-Bao, HU Liang, MENG Qing-Yin. Responses of nitrogen and phosphorus resorption from leaves and branches to long-term nitrogen deposition in a Chinese fir plantation. Chinese Journal of Plant Ecology, 2018, 42(9): 926-937. DOI: 10.17521/cjpe.2018.0167

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图/表 9

表1 各处理样地的主要林分特征和土壤理化性质本底值

Table 1 Background values of the stand and soil physicochemical properties in the nitrogen addition plots

处理 Treatment	林分特征 Stand characteristics				土壤理化性质 Soil physicochemical properties
处理 Treatment	林龄 Forest age (a)	密度 Density (No.·hm^-2)	平均胸径 Mean DBH (cm)	平均树高 Mean tree height (m)	pH	有机碳 Organic carbon (g·kg^-1)	全氮 Total N (g·kg^-1)	全磷 Total P (g·kg^-1)
N0	12	1 717	16.1	11.8	4.59	19.23	0.86	0.22
N1	12	1 633	16.0	12.2	4.76	17.31	0.68	0.13
N2	12	1 683	16.3	12.2	4.65	18.88	0.80	0.17
N3	12	1 625	16.0	12.1	4.71	18.14	0.81	0.14

表2 基于双因素方差分析得到的组分、氮沉降水平及其交互作用对杉木叶和枝的生态化学计量的影响(F值)

Table 2 Results (F values) of two-way ANOVA on the effects of components, nitrogen deposition treatment level and their interaction the ecological stoichiometry in leaves and branches of Chinese fir

因子 Factor	F (p)值 F (p) value
因子 Factor	C	N	P	C:N	C:P	N:P
叶组分 Leaf components	1.173 (0.336)	20.401 (<0.001)	24.425 (<0.001)	13.832 (<0.001)	24.072 (<0.001)	5.728 (0.003)
氮沉降 N deposition	0.573 (0.637)	3.272 (0.034)	1.246 (0.309)	3.158 (0.038)	0.853 (0.475)	0.631 (0.601)
叶组分×氮沉降 Leaf components × N deposition	0.280 (0.976)	0.893 (0.543)	0.865 (0.565)	0.537 (0.837)	0.963 (0.487)	0.575 (0.807)
枝组分 Branch components	1.191 (0.329)	9.407 (<0.001)	19.685 (<0.001)	6.534 (0.001)	6.010 (0.002)	8.176 (<0.001)
氮沉降 N deposition	0.165 (0.919)	1.782 (0.170)	2.130 (0.116)	1.229 (0.315)	1.063 (0.378)	5.123 (0.005)
枝组分×氮沉降 Branch components × N deposition	0.398 (0.927)	0.608 (0.781)	0.365 (0.943)	0.429 (0.910)	0.371 (0.940)	0.442 (0.902)

图2 杉木叶和枝的测试指标(C、N、P、C:N、C:P、N:P)氮沉降处理平均值与N0处理平均值的百分比(N0/N0 = 100%)。A, 一年生叶。B, 二年生叶。C, 衰老叶。D, 一年生枝。E, 二年生枝。F, 衰老枝。N0、N1、N2、N3, 施氮量分别为0、60、120、240 kg·hm-2·a-1。

Fig. 2 Mean percentage changes of measured parameters (C, N, P, C:N, C:P, N:P) of Chinese fir leaf and branch after N deposition treatment for 14 years relative to these values under control (N0) (N0/N0 = 100%). A, One-year old leaf. B, Two-year old leaf. C, Senesced leaf. D, One-year old branch. E, Two-year old branch. F, Senesced branch. N0, N1, N2, N3, nitrogen addition 0, 60, 120, 240 kg·hm-2·a-1, respectively.

图3 长期氮沉降条件下杉木叶和枝的N、P再吸收率(平均值+标准误差)。黑色柱子表示一年生叶/枝与衰老叶/枝之间的养分再吸收率; 浅灰柱子表示二年生叶/枝与衰老叶/枝之间的养分再吸收率; 白色柱子表示一年生叶/枝与二年生叶/枝之间的养分再吸收率。N0、N1、N2、N3, 施氮量分别为0、60、120、240 kg·hm-2·a-1。不同小写字母表示不同氮沉降水平下达到显著差异水平(p < 0.05)。

Fig. 3 Effects of long-term nitrogen deposition on N and P resorption efficiency in leaves and branches of Chinese fir (mean + SE). The black column indicates the ratio of the nutrient resorption efficiency between one-year old leaves/branches and senescent leaves/branches; light gray column indicates the ratio of the nutrient resorption efficiency between two-year old leaves/branches and senescent leaves/branches; white column indicates the ratio of the nutrient resorption efficiency between one-year old leaves/branches and two-year-old leaves. N0, N1, N2, N3, nitrogen addition 0, 60, 120, 240 kg·hm-2·a-1, respectively. Different lowercase letters indicate significant differences under different nitrogen addition treatment levels (p < 0.05).

图4 长期氮沉降条件下杉木叶和枝REN与REP, N:P与REN:REP的线性关系。

Fig. 4 Linear relationship between N resorption efficiency (REN) and P resorption efficiency (REP), N:P and N resorption efficiency to P resorption efficiency ratio (REN:REP) of leaf and branch under long-term nitrogen deposition treatments. Data included treatments for 14 years.

图5 2010年成熟绿叶和衰老叶的N、P含量(平均值+标准误差)。N1、N2、N3, 施氮量分别为0、60、120、240 kg·hm-2·a-1。

Fig. 5 N and P content of mature living and senescent leaves in 2010 (mean + SE). N0, N1, N2, N3, nitrogen addition 0, 60, 120, 240 kg·hm-2·a-1, respectively.

图6 2010和2017年的叶N、P再吸收率(平均值+标准误差)。N1、N2、N3, 施氮量分别为0、60、120、240 kg·hm-2·a-1。

Fig. 6 N and P resorption efficiency of leaves in 2010 and 2017 (mean + SE). N0, N1, N2, N3, nitrogen addition 0, 60, 120, 240 kg·hm-2·a-1, respectively.

表3 2010和2017年的叶和枝N再吸收率与P再吸收率的比值

Table 3 The ratio of N resorption efficiency to P resorption efficiency (REN:REP) of leaves and branches in 2010 and 2017

处理 Treatment	2010	2017
处理 Treatment	成熟-衰叶Mature- senescent leaf	一-衰叶 One-senesced leaf	二-衰叶 Two-senesced leaf	一-衰枝 One-senesced branch	二-衰枝 Two-senesced branch
N0	0.89	0.69	0.37	0.69	0.75
N1	0.82	0.43	0.40	1.40	1.59
N2	0.87	0.63	0.34	1.06	0.67
N3	0.88	0.70	0.35	0.95	0.76

图1 长期氮沉降条件下杉木叶和枝的C、N、P含量及化学计量(平均值+标准误差)。不同小写字母表示相同龄级叶或枝不同氮沉降水平下差异显著(p < 0.05)。N0、N1、N2、N3, 施氮量分别为0、60、120、240 kg·hm-2·a-1。

Fig. 1 C, N, P content and stoichiometry of leaves and branches of Chinese fir under long-term nitrogen deposition (mean + SE). Different lowercase letters indicate significant differences for the same life span leaf or branch under different nitrogen deposition treatment levels (p < 0.05). N0, N1, N2, N3, nitrogen addition 0, 60, 120, 240 kg·hm-2·a-1, respectively.

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长期氮沉降对杉木人工林叶、枝氮磷养分再吸收的影响

Responses of nitrogen and phosphorus resorption from leaves and branches to long-term nitrogen deposition in a Chinese fir plantation

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