Effects of nitrogen addition and planting density on the growth and biological nitrogen fixation of Lespedeza davurica

doi:10.17521/cjpe.2020.0185

Abstract

Abstract:

Aims Nitrogen (N) supply and planting density are two important factors influencing plant growth. Legumes are important to ecosystem N input because of their capacity of biological N₂ fixation (BNF). Increasing atmospheric N deposition may promote the growth of leguminous plants, but it may also inhibit strongly their BNF capacity. Planting density can also influence the growth and BNF capacity of legumes due to intraspecific competition. However, few studies up to date have focused on the effects of N fertilization and planting density on the growth and BNF capacity of legumes. In this study, we aimed to explore the potentially interactive effects of N fertilization and planting density on the growth and BNF capacity of Lespedeza davurica, a leguminous plant species which is widely distributed throughout the northern China grasslands.
Methods A pot experiment was conducted in a greenhouse. The experiment contained four levels of N addition (0, 5, 10, 20 g·m^-2·a^-1) by NH₄NO₃fertilizer and three levels of planting densities (1, 3, 6 Ind.·pot^-1, i.e. 32, 96, 192 Ind.·m^-2).
Important findings 1) Our results showed that both N addition and planting density could impact the biomass production ofL. davurica. N addition increased plant leaf carbon (C) and N contents and leaf-level net photosynthetic rate. Besides, N addition also stimulated the plant growth at both pot and individual levels, and the yield reached maximum at N addition of 10 g·m^-2·a^-1. Increasing planting density decreased leaf C and N contents, leaf-level net photosynthetic rate, and individual growth, but increased total biomass in each pot. 2) Nitrogen addition reduced the capacity of BNF of L. davurica, while increasing planting density could weaken this suppression effect to some extent. The combination of N addition of 10 g·m^-2·a^-1 and planting density of 3 Ind.·pot^-1 or N addition of 10 g·m^-2·a^-1 and planting density of 6 Ind.·pot^-1 could maximize the effects of N application on individual yield and the effects of increasing planting density on the alleviation of BNF suppression. Nitrogen addition suppressed the BNF of L. davurica through reducing plant investment to nodulation and nodule biomass production. The intraspecific competition and resource limitation caused by increasing planting density led to improvement in the investment to nodulation and nodule growth. 3) Structural equation model analyses showed that N addition and planting density combined explained variations in the plant biomass and nodule production either directly or indirectly by 64% and 42%, respectively. The results indicate that it is important to optimize the amount of fertilizer application and appropriate planting density when considering plantation and management of artificial and degraded grasslands.

Key words: N addition, planting density, biomass, nodule biomass, investment to nodulation, biological nitrogen fixation

WANG Yin-Liu, GENG Qian-Qian, HUANG Jian-Hui, WANG Chang-Hui, LI Lei, HASI Muqier, NIU Guo-Xiang. Effects of nitrogen addition and planting density on the growth and biological nitrogen fixation of Lespedeza davurica[J]. Chin J Plant Ecol, 2021, 45(1): 13-22.

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URL: https://www.plant-ecology.com/EN/10.17521/cjpe.2020.0185

https://www.plant-ecology.com/EN/Y2021/V45/I1/13

Figures/Tables 5

Table 1 Growth indexes of Lespedeza davurica under different levels of nitrogen addition and planting density treatments (mean ± SE)

处理 Treatment	水平 Level	株高 Height (cm)	总生物量 Total biomass (g)	总地上生物量 Total aboveground biomass (g)	总地下生物量 Total belowground biomass (g)	平均地上生物量 Average aboveground biomass (g·Ind.^-1)	平均地下生物量 Average belowground biomass (g·Ind.^-1)	根冠比 Root:shoot ratio	叶片C含量 Leaf C concentration (%)
施氮 N addition (g·m^-2·a^-1)	0	58.69 ± 2.14 ^ns	11.79 ± 1.02 ^b	8.94 ± 0.72 ^c	2.84 ± 0.32 ^b	3.61 ± 0.47 ^b	1.06 ± 0.13 ^b	0.31 ± 0.02 ^b	45.26 ± 0.34 ^b
	5	58.49 ± 1.90 ^ns	15.43 ± 1.02 ^a	11.24 ± 0.64 ^bc	4.20 ± 0.39 ^a	4.93 ± 0.76 ^a	1.67 ± 0.22 ^a	0.36 ± 0.02 ^a	45.99 ± 0.35 ^a
	10	62.56 ± 2.43 ^ns	15.76 ± 1.04 ^a	11.99 ± 0.77 ^a	3.76 ± 0.32 ^a	5.20 ± 0.80 ^a	1.60 ± 0.25 ^a	0.31 ± 0.02 ^b	45.98 ± 0.32 ^a
	20	63.32 ± 2.61 ^ns	15.37 ± 1.02 ^a	11.52 ± 0.7 ^ab	3.86 ± 0.33 ^a	4.91 ± 0.67 ^a	1.51 ± 0.17 ^a	0.33 ± 0.01 ^ab	45.95 ± 0.26 ^a
密度 Planting density (Ind.·pot^-1)	1	77.53 ± 4.43 ^a	10.01 ± 0.62 ^b	7.80 ± 0.45 ^b	2.20 ± 0.19 ^b	7.81 ± 0.45 ^a	2.20 ± 0.19 ^a	0.28 ± 0.01 ^b	46.29 ± 0.09 ^a
	3	61.56 ± 1.60 ^b	16.54 ± 0.62 ^a	12.20 ± 0.46 ^a	4.34 ± 0.19 ^a	4.07 ± 0.15 ^b	1.45 ± 0.06 ^b	0.36 ± 0.01 ^a	46.69 ± 0.24 ^a
	6	57.53 ± 1.42 ^b	17.50 ± 0.55 ^a	12.94 ± 0.38 ^a	4.56 ± 0.25 ^a	2.16 ± 0.06 ^c	0.76 ± 0.04 ^c	0.35 ± 0.02 ^a	44.41 ± 0.13 ^b
N × density		**	***	***	***	***	***	***	ns

Fig. 1 Changes in leaf nitrogen (N) concentration (A) and net photosynthetic rate (B) of Lespedeza davurica under different treatments (mean ± SE). Different lowercase letters indicate significant difference among N addition treatments under the same density at p < 0.05 level, while different uppercase letters indicate significant difference among density treatments at p < 0.05 level. ns, p > 0.05; *, p < 0.05; **, p < 0.01; ***, p < 0.001.

Fig. 2 Relative neighbor effect index under different treatments (mean ± SE). Different lowercase letters indicate significant difference among density treatments under the same N addition rate at p < 0.05 level. ns, p > 0.05; *, p < 0.05; **, p < 0.01; ***, p < 0.001.

Fig. 3 Responses of the nodule biomass response ratio (RR)(A) and investment to nodulation (B) under different treatments (mean ± SE). In Fig. A, response ratio values with error bars not overlapping RR = 1 (horizontal dotted line), indicate significant difference between treatment and control (p < 0.05). *, p < 0.05; **, p < 0.01; ***, p < 0.001.

Fig. 4 Direct and indirect impacts of N addition rate and planting density on the root nodulation and growth of Lespedeza davurica. Lines indicate a significant effect (p < 0.05). Line thickness indicates relative effect size. Black lines represent positive effects, while gray lines indicate negative effects. R 2 indicates variation that can be explained. Pn, net photosynthetic rate.

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