不同种源刨花楠苗木生长与主要功能性状对氮添加的响应

doi:10.17521/cjpe.2022.0301

植物生态学报 ›› 2023, Vol. 47 ›› Issue (12): 1693-1707.DOI: 10.17521/cjpe.2022.0301

不同种源刨花楠苗木生长与主要功能性状对氮添加的响应

安凡¹, 李宝银¹^,²^,^*(), 钟全林¹^,², 程栋梁¹^,², 徐朝斌¹, 邹宇星¹, 张雪¹, 邓兴宇¹, 林秋燕¹

¹福建师范大学地理科学学院, 福州 350007
²湿润亚热带山地生态国家重点实验室培育基地, 福建师范大学福建省植物生理生态重点实验室, 福州 350007

收稿日期:2022-07-20 接受日期:2023-01-31 出版日期:2023-12-20 发布日期:2023-02-03
通讯作者: *(liby@fjnu.edu.cn)
基金资助:
国家自然科学基金(31971643);国家自然科学基金(32071555);福建省科技厅高校产学合作项目(2019N5009和2020N5008(2019N5009);福建省科技厅高校产学合作项目(2019N5009和2020N5008(2020N5008)

Nitrogen addition affects growth and functional traits of Machilus pauhoi seedlings from different provenances

AN Fan¹, LI Bao-Yin¹^,²^,^*(), ZHONG Quan-Lin¹^,², CHENG Dong-Liang¹^,², XU Chao-Bin¹, ZOU Yu-Xing¹, ZHANG Xue¹, DENG Xing-Yu¹, LIN Qiu-Yan¹

¹College of Geographical Sciences, Fujian Normal University, Fuzhou 350007, China
²State Key Laboratory Breeding Base of Humid Subtropical Mountain Ecology, Fujian Province Key Laboratory of Plant Ecophysiology, Fujian Normal University, Fuzhou 350007, China

Received:2022-07-20 Accepted:2023-01-31 Online:2023-12-20 Published:2023-02-03
Contact: *(liby@fjnu.edu.cn)
Supported by:
National Natural Science Foundation of China(31971643);National Natural Science Foundation of China(32071555);Industry-University Cooperation Project of Fujian Science and Technology Department(2019N5009);Industry-University Cooperation Project of Fujian Science and Technology Department(2020N5008)

摘要/Abstract

摘要：

氮是影响植物生长的重要营养物质元素, 生物量分配模式和功能性状反映植物在不同环境中的适应性策略, 揭示不同种源个体对氮沉降的生态响应, 有助于阐明植物对氮沉降的适应机制。该研究以刨花楠(也称刨花润楠, Machilus pauhoi)典型分布区福建万木林、福建茫荡山、江西遂川、浙江建德、湖南茶陵、江西安福6个种源地的刨花楠苗木为对象, 通过设置施肥和不施肥处理, 探讨氮添加、种源及其交互作用对刨花楠苗木生物量分配与功能性状的影响。结果表明: 1)氮添加会显著促进刨花楠苗木的生物量累积及其相对生长速率, 建德种源苗木在施氮处理后长势最好; 氮添加对各种源苗木生物量分配的影响表现不同, 添加氮会显著增加建德和安福种源苗木叶质量分数及万木林和安福种源苗木的叶茎比。2)氮添加对各种源苗木叶性状的影响存在差异; 施氮会显著降低建德、茶陵和安福种源苗木的比叶面积, 增加茶陵种源苗木的叶面积及叶片氮磷比。3)各种源苗木细根主要功能性状对氮添加的响应也存在差异; 施氮会显著降低建德种源苗木的细根平均直径及遂川、建德和安福种源苗木的根组织密度, 但却会增加万木林种源苗木根组织密度、茫荡山和建德种源苗木的比根长及遂川和建德种源苗木的细根比表面积; 氮添加显著增加安福种源苗木的细根氮含量, 降低万木林、茫荡山和遂川种源苗木的细根磷含量, 并使各种源苗木细根氮磷比均显著高于对照。4)不同种源间刨花楠苗木细根和与生物量分配相关的表型可塑性相对高于叶片表型可塑性, 其中建德种源苗木的表型可塑性最高, 遂川种源苗木表型可塑性最低。5)结构方程模型表明, 种源与氮添加对刨花楠苗木生长影响显著, 氮添加通过影响叶茎比和叶片氮磷比进而对生长产生影响, 种源则通过影响其比叶面积与细根比表面积进而影响其生长。不同种源间刨花楠苗木生长、生物量分配及主要功能性状对氮添加的生态适应策略具有差异性, 其中建德种源苗木更适应外界环境的变化, 这是其长期适应种源地环境的结果。研究结果可为我国受氮沉降影响的亚热带区域开展刨花楠优良种源选择等提供理论依据。

关键词: 种源, 氮添加, 功能性状, 生长, 刨花楠

Abstract:

Aims Nitrogen (N) is an important nutrient element affecting plant growth, biomass allocation and functional trait strategies. Revealing the ecological responses of different provenances to N deposition can help elucidate the adaptation mechanisms of plant species.
Methods In this study, Machilus pauhoi seedlings from 6 provenances in Wanmulin, Fujian Province (WML), Mangdang Mountain, Fujian Province (MDS), Suichuan, Jiangxi Province (SC), Jiande, Zhejiang Province (JD), Chaling, Hunan Province (CL) and Anfu, Jiangxi Province (AF) were selected as our study objects. By setting two treatments of fertilization and no fertilization, we investigated the effects of N addition, provenance and their interactions on biomass allocation and functional traits of M. pauhoi seedlings.
Important findings 1) Nitrogen addition significantly promoted total biomass accumulation and the relative growth rate of M. pauhoi seedlings, with the greatest response observed for JD seedlings. However, the effect of N addition on biomass allocation varied by provenance, with JD and AF seeding showing increased leaf mass fraction, and WML and AF seedlings showing increased leaf stem ratio. 2) The effects of N addition on leaf traits of seedlings also varied by provenance. N application significantly reduced the specific leaf area of JD, CL and AF seedlings, but increased the leaf area and leaf N:phosphorus (P) of CL seedling. 3) The response of major functional traits of fine roots to N addition also varied by provenance. N addition significantly decreased the average root diameter and root tissue density of SC, JD and AF seedlings, but increased the root tissue density of WML seedling, the specific root length of MDS and JD seedlings, and the specific surface area of SC and JD seedlings. N addition significantly increased the fine root N content of AF seedling, but decreased the fine root P content of WML, MDS and SC seedlings, and the fine root N:P of all provenances were significantly higher in fertilization treatments than in the control. 4) We observed higher phenotypic plasticity in fine root and biomass-related phenotypes than in leaf traits among different provenances, with the highest phenotypic plasticity in the JD seedling and the lowest phenotypic plasticity in the SC seedling. 5) Structural equation modeling showed that N addition and provenance had significant effects on the growth of M. pauhoi seedlings, with N addition influencing growth by affecting leaf stem ratio and leaf N:P, and provenance influencing growth by affecting specific leaf area and fine root surface area. The growth, biomass and functional traits of M. pauhoi seedlings from different provenances have different ecological adaptation strategies to N addition. Among them, JD seedlings is more adaptable to changes in the external environment, which may account for its long-term adaptation to the provenance environment. Our results provide a theoretical basis for the selection of optimal provenance of M. pauhoi in subtropical regions of China that affected by N deposition.

Key words: provenances, nitrogen addition, functional trait, growth, Machilus pauhoi

安凡, 李宝银, 钟全林, 程栋梁, 徐朝斌, 邹宇星, 张雪, 邓兴宇, 林秋燕. 不同种源刨花楠苗木生长与主要功能性状对氮添加的响应. 植物生态学报, 2023, 47(12): 1693-1707. DOI: 10.17521/cjpe.2022.0301

AN Fan, LI Bao-Yin, ZHONG Quan-Lin, CHENG Dong-Liang, XU Chao-Bin, ZOU Yu-Xing, ZHANG Xue, DENG Xing-Yu, LIN Qiu-Yan. Nitrogen addition affects growth and functional traits of Machilus pauhoi seedlings from different provenances. Chinese Journal of Plant Ecology, 2023, 47(12): 1693-1707. DOI: 10.17521/cjpe.2022.0301

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2022.0301

https://www.plant-ecology.com/CN/Y2023/V47/I12/1693

图/表 10

表1 刨花楠种源地地理环境概况

Table 1 Geographical environment characteristics of Machilus pauhoi provenances

种源地 Provenance	纬度 Latitude (N)	经度 Longitude (E)	年平均气温 Annual mean temperature (℃)	年降水量 Annual precipitation (mm)
WML	27.05°	118.12°	18.7	1 663
MDS	26.41°	118.09°	19.3	1 663
SC	26.21°	114.23°	17.5	1 421
JD	29.25°	119.07°	17.4	1 712
CL	26.39°	113.46°	20.0	1 410
AF	26.41°	118.07°	19.3	1 663

表2 氮添加及种源对刨花楠苗木生长和功能性状的双因素方差分析(F值)

Table 2 Two-way ANOVA of the effects of nitrogen (N) addition and provenance on the growth and functional traits of Machilus pauhoi seedlings (F value)

性状 Trait	种源 Provenance	氮添加 N addition	种源×氮添加 Provenance × N addition
RGR	8.30^**	30.38^**	0.51
RHI	10.48^**	39.31^**	2.86
LMF	1.85	9.22^**	1.06
RMF	7.21^*	2.07	0.95
RSR	4.74^*	1.56	1.39
LSR	3.37^**	16.46^**	1.01
LNC	0.97	0.49	0.57
LPC	1.43^*	1.69	1.01
LN/P	0.85	2.41^*	0.86
LMI	1.19	0.09	0.76
SLA	5.46^**	21.76^**	3.77^**
LA	5.26^*	3.23	2.06
LDMC	21.84^*	0.48	0.52
AD	2.38^*	5.90^*	0.97
SRL	24.04^**	1.06	2.07
SRA	11.15^**	2.73	1.19
RTD	0.69	1.29	1.83
RNC	0.42	0.43	1.76
RPC	1.39	31.70^**	2.91^*
RN/P	1.31	33.31^**	1.16

图1 氮添加对不同种源刨花楠苗木相对生长速率的影响(平均值±标准误)。CT, 对照; +N, 氮添加。AF, 江西安福; CL, 湖南茶陵; JD, 浙江建德; MDS, 福建茫荡山; SC, 江西遂川; WML, 福建万木林。不同小写字母表示同一处理不同种源间的差异显著(p < 0.05), 不同大写字母表示同一种源不同处理间的差异显著(p < 0.05)。

Fig. 1 Effect of nitrogen addition on the relative growth rate of Machilus pauhoi seedlings from different provenances (mean ± SE). CT, control treatment; +N, nitrogen addition. AF, Anfu, Jiangxi Province; CL, Chaling, Hunan Province; JD, Jiande, Zhejiang Province; MDS, Mangdang Mountain, Fujian Province; SC, Suichuan, Jiangxi Province; WML, Wanmulin, Fujian Province. Different lowercase letters indicate significant differences between different provenances of the same treatment, and different uppercase letters indicate significant differences between different treatments of the same provenances (p < 0.05).

图2 氮添加对刨花楠苗木生物量分配的影响(平均值±标准误)。CT, 对照; +N, 氮添加。AF, 江西安福; CL, 湖南茶陵; JD, 浙江建德; MDS, 福建茫荡山; SC, 江西遂川; WML, 福建万木林。不同小写字母表示同一处理不同种源间的差异显著(p < 0.05), 不同大写字母表示同一种源不同处理间的差异显著(p < 0.05)。

Fig. 2 Effect of nitrogen addition on the biomass allocation in different provenances of Machilus pauhoi seedlings (mean ± SE). CT, control treatment; +N, nitrogen addition. AF, Anfu, Jiangxi Province; CL, Chaling, Hunan Province; JD, Jiande, Zhejiang Province; MDS, Mangdang Mountain, Fujian Province; SC, Suichuan, Jiangxi Province; WML, Wanmulin, Fujian Province. Different lowercase letters indicate significant differences between different provenances of the same treatment, and different uppercase letters indicate significant differences between different treatments of the same provenances (p < 0.05).

图3 氮添加对刨花楠苗木叶片表型性状的影响(平均值±标准误)。CT, 对照; +N, 氮添加。AF, 江西安福; CL, 湖南茶陵; JD, 浙江建德; MDS, 福建茫荡山; SC, 江西遂川; WML, 福建万木林。不同小写字母表示同一处理不同种源间的差异显著(p < 0.05), 不同大写字母表示同一种源不同处理间的差异显著(p < 0.05)。

Fig. 3 Effect of nitrogen addition on leaf traits of Machilus pauhoi seedling from different provenances (mean ± SE). CT, control treatment; +N, nitrogen addition. AF, Anfu, Jiangxi Province; CL, Chaling, Hunan Province; JD, Jiande, Zhejiang Province; MDS, Mangdang Mountain, Fujian Province; SC, Suichuan, Jiangxi Province; WML, Wanmulin, Fujian Province. Different lowercase letters indicate significant differences between different provenances of the same treatment, and different uppercase letters indicate significant differences between different treatments of the same provenances (p < 0.05).

图4 氮添加对刨花楠苗木细根性状的影响(平均值±标准误)。CT, 对照; +N, 氮添加。AF, 江西安福; CL, 湖南茶陵; JD, 浙江建德; MDS, 福建茫荡山; SC, 江西遂川; WML, 福建万木林。不同小写字母表示同一处理不同种源间的差异显著(p < 0.05), 不同大写字母表示同一种源不同处理间的差异显著(p < 0.05)。

Fig. 4 Effect of nitrogen addition on fine root traits of Machilus pauhoi seedling from different provenances (mean ± SE). CT, control treatment; +N, nitrogen addition. AF, Anfu, Jiangxi Province; CL, Chaling, Hunan Province; JD, Jiande, Zhejiang Province; MDS, Mangdang Mountain, Fujian Province; SC, Suichuan, Jiangxi Province; WML, Wanmulin, Fujian Province. Different lowercase letters indicate significant differences between different provenances of the same treatment, and different uppercase letters indicate significant differences between different treatments of the same provenances (p < 0.05).

表3 氮添加对刨花楠苗木叶片和细根氮(N)、磷(P)化学计量特征的影响(平均值±标准误)

Table 3 Effect of nitrogen addition on the stoichiometric ratio of nitrogen (N) and phosphorus (P) in leaves and fine roots of Machilus pauhoi seedling from different provenances (mean ± SE)

种源 Provenance	处理 Treatment	叶片 Leaf			细根 Fine root
种源 Provenance	处理 Treatment	LNC (mg·g^-1)	LPC (mg·g^-1)	N:P	RNC (mg·g^-1)	RPC (mg·g^-1)	N:P
WML	CT	15.77 ± 0.97^Aa	1.47 ± 0.09^Aa	10.73 ± 0.30^Aa	10.22 ± 0.58^Aa	3.00 ± 0.27^Aa	3.62 ± 0.25^Aa
WML	+N	15.69 ± 0.82^Aa	1.39 ± 0.06^Aa	12.27 ± 0.32^Aab	10.41 ± 0.37^Aab	1.82 ± 0.17^Ba	5.97 ± 0.52^Bc
MDS	CT	14.99 ± 1.10^Aa	1.28 ± 0.08^Aa	12.01 ± 1.41^Aa	10.64 ± 0.66^Aa	2.69 ± 0.43^Aa	4.40 ± 0.62^Aa
MDS	+N	13.53 ± 0.84^Aa	1.15 ± 0.09^Aab	11.99 ± 1.01^Aab	9.45 ± 0.37^Ab	1.31 ± 0.12^Bb	7.46 ± 0.61^Bbc
SC	CT	13.96 ± 0.59^Aa	1.22 ± 0.11^Aa	12.00 ± 0.83^Aa	10.12 ± 0.31^Aa	2.44 ± 0.21^Aa	4.33 ± 0.30^Aa
SC	+N	14.09 ± 1.01^Aa	1.22 ± 0.09^Aab	11.54 ± 0.43^Ab	9.99 ± 0.27^Aab	1.58 ± 0.13^Bab	6.71 ± 0.59^Bbc
JD	CT	14.65 ± 1.01^Aa	1.43 ± 0.10^Aa	11.94 ± 0.47^Aa	9.72 ± 0.64^Aa	1.95 ± 0.17^Aa	5.09 ± 0.22^Aa
JD	+N	16.12 ± 0.70^Aa	1.24 ± 0.10^Aab	11.56 ± 0.55^Ab	10.93 ± 0.61^Aa	1.67 ± 0.34^Aab	7.85 ± 0.81^Bbc
CL	CT	14.84 ± 0.60^Aa	1.32 ± 0.12^Aa	11.36 ± 0.74^Ba	10.49 ± 0.61^Aa	2.25 ± 0.43^Aa	4.91 ± 0.66^Aa
CL	+N	12.64 ± 0.79^Aa	0.89 ± 0.16^Ab	14.97 ± 2.25^Aa	11.08 ± 0.76^Aa	1.21 ± 1.14^Ab	11.50 ± 4.43^Ba
AF	CT	15.11 ± 1.17^Aa	1.41 ± 0.15^Aa	11.13 ± 0.79^Aa	9.50 ± 0.57^Aa	2.07 ± 0.17^Aa	4.71 ± 0.25^Aa
AF	+N	14.30 ± 0.99^Aa	1.25 ± 0.12^Aab	11.90 ± 0.82^Ab	10.54 ± 0.23^Ba	1.62 ± 0.21^Aab	8.27 ± 1.27^Bb

图5 不同种源刨花楠苗木的表型可塑性指数。AF, 江西安福; CL, 湖南茶陵; JD, 浙江建德; MDS, 福建茫荡山; SC, 江西遂川; WML, 福建万木林。AD, 细根平均直径; LA, 叶面积; LDMC, 叶干物质含量; LMF, 叶质量分数; LMI, 叶形态指数; LNC, 叶片氮含量; LN/P, 叶片氮磷比; LPC, 叶片磷含量; LSR, 叶茎比; RMF, 根质量分数; RNC, 细根氮含量; RN/P, 细根氮磷比; RPC, 细根磷含量; RSR, 根冠比; RTD, 根组织密度; SLA, 比叶面积; SRA, 细根比表面积; SRL, 比根长。

Fig. 5 Index of phenotypic plasticity of Machilus pauhoi seedings of different provenances. AF, Anfu, Jiangxi Province; CL, Chaling, Hunan Province; JD, Jiande, Zhejiang Province; MDS, Mangdang Mountain, Fujian Province; SC, Suichuan, Jiangxi Province; WML, Wanmulin, Fujian Province. AD, average fine root diameter; LA, leaf area; LDMC, leaf dry matter content; LMF, leaf mass fraction; LMI, leaf morphology index; LNC, leaf nitrogen (N) content; LN/P, leaf N:phosphorus (P); LPC, leaf P content; LSR, leaf stem biomass ratio; RMF, root mass fraction; RNC, root N content; RN/P, root N:P; RPC, root P content; RSR, root shoot biomass ratio; RTD, root tissue density; SLA, specific leaf area; SRA, specific fine root area; SRL, specific root length.

表4 不同种源刨花楠苗木生长与功能性状的逐步回归分析

Table 4 Stepwise regression analysis of growth and leaf functional traits of Machilus pauhoi seedling from different provenances

指标 Index	回归方程 Regression equation	标准化回归系数 Standardized regression coeffcient	R²
相对生长速率 RGR	Y = -0.25 + 0.53N + 0.014Lng + 0.005MAT	B_N = 0.47, B_Lng = 0.302, B_MAT = -0.21	0.42^*
比叶面积 SLA	Y = 159.153 - 11.2N + 9.97Lng - 3.79Pr - 2.32Lat	B_N = -0.34, B_Lng = 0.67, B_Pr = -0.48, B_Lat = -0.30	0.26^*
叶面积 LA	Y = 6.30 + 0.89Pr + 0.66MAT	B_Pr = 0.35, B_MAT = 0.32	0.14^**
叶干物质含量 LDMC	Y = 4.59 + 0.14Pr - 0.11Lng - 0.5MAT + 0.013MAP + 0.001Lat	B_Pr = 1.07, B_Lng = -0.43, B_MAT = 1.22, B_MAP = 4.59, B_Lat = -3.78	0.58^*
比根长 SRL	Y = 4.43 + 0.13Pr - 0.12Lng + 0.042Lat	B_Pr = 0.68, B_Lng = -0.36, B_Lat = 0.25	0.53^*
细根磷含量 RPC	Y = 3.56 - 0.81N - 0.10Pr	B_N = -0.51, B_Pr = -0.22	0.35^*
细根氮磷比 Root N:P	Y = 0.41 + 3.06N + 0.29Pr	B_N = -0.51, B_Pr = -0.22	0.35^*

图6 氮添加和种源对刨花楠苗木生长的直接和间接效应。图中橙色实线箭头表示负向路径(p < 0.05), 蓝色实线箭头表示正向路径(p < 0.05), 灰色虚线箭头表示非显著路径(p > 0.05), 箭头宽度表示因果关系的强弱。线条上的与箭头相邻的数字是标准化的路径系数。R2表示模型解释的总变异量。*, p < 0.05; **, p < 0.01。GFI, 拟合优度指数; LN/P, 叶片氮磷比; LSR, 叶茎比; RGR, 生物量相对生长速率; RMSEA, 逼近均方误差; SLA, 比叶面积; SRA, 细根比表面积; SRMR, 标准化均方根残差。

Fig. 6 Direct and indirect effects of nitrogen (N) addition and seed source on the growth of Machilus pauhoi seedlings. Solid orange arrows in the figure indicate negative paths (p < 0.05), solid blue arrows indicate positive paths (p < 0.05), and dashed gray arrows indicate non-significant paths (p > 0.05). The width of the arrow indicates the strength of the causal relationship. Numbers beside the line are the standardized path coefficients. R2 indicates the total variance explained by the model. GFI, goodness of fit index; LN/P, leaf N:P; LSR, leaf stem biomass ratio; RGR, relevant growth rate; RMSEA, root mean square error of approximation; SRA, specific fine root area; SLA, specific leaf area; SRMR, standardized root mean square residual. *, p < 0.05; **, p < 0.01.

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不同种源刨花楠苗木生长与主要功能性状对氮添加的响应

Nitrogen addition affects growth and functional traits of Machilus pauhoi seedlings from different provenances

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