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

doi:10.17521/cjpe.2022.0301

Abstract

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

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[J]. Chin J Plant Ecol, 2023, 47(12): 1693-1707.

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URL: https://www.plant-ecology.com/EN/10.17521/cjpe.2022.0301

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

Figures/Tables 10

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

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

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).

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).

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).

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).

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

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.

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^*

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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