恢复方式对东北东部森林土壤碳氮磷计量特征的影响

doi:10.17521/cjpe.2023.0234

摘要/Abstract

摘要：

土壤生态化学计量是土壤肥力和植物养分状况的重要指标, 探索森林的恢复方式对土壤生态化学计量特征的影响, 可为准确评价生态系统物质循环过程对干扰和恢复的响应提供理论依据和数据支撑。该研究比较测定了东北东部山区人工恢复的3种针阔混交林和天然恢复的4种落叶阔叶林的土壤、凋落物和枯落物的碳(C)、氮(N)和磷(P)含量, 以及土壤pH、土壤密度等理化性质。结果显示: 人工针阔混交林和天然落叶阔叶林的土壤C、N和P含量均随着土壤深度的增加而降低。各林分类型的O (有机质)层土壤C、N和P含量的波动范围分别为53.78-90.59、5.02-7.83和0.75-0.91 g·kg^-1。人工针阔混交林O层土壤的C和N含量显著低于天然落叶阔叶林, 而人工针阔混交林A (腐殖质)和B (淀积)层土壤的C、N和P含量均高于天然落叶阔叶林。人工针阔混交林O层土壤的C密度显著低于天然阔叶林。土壤C:N、C:P和N:P的波动范围为10.08-12.53、43.97-135.52和4.56-11.64; O层土壤的C:N在两种恢复方式的森林间无显著差异, 人工针阔混交林O和A层土壤的C:P和N:P显著低于天然落叶阔叶林。土壤各层次的C和N含量均存在显著的正相关关系(R²范围0.40-0.76)。除C:N以外, 恢复方式、土壤发生层及其二者交互作用对土壤C、N、P含量、密度和计量比均存在显著影响。土壤密度和凋落物C含量显著影响土壤C、N和P含量。这些结果表明, 通过人工恢复方式增加针叶树种的比重, 使表层土壤的C、N含量降低, 导致表层土壤的碳固持量减弱; 但是对C:N无显著影响, 表现出相对稳定的土壤C、N计量特征。

关键词: C:N:P, 生态化学计量, 森林恢复, 养分循环, 土壤碳固持

Abstract:

Aims Soil ecological stoichiometry is an important indicator of soil fertility and plant nutrient status. Investigating impacts of restoration approaches on forest soil ecological stoichiometry characteristics provides a theoretical foundation and data support for accurately assessing responses of the ecosystem material cycling processes to disturbance and restoration efforts.
Methods This study compared the forest soil stoichiometry from two forest restoration approaches, i.e., artificial (RG) and natural regenerations (TR) that included three coniferous or broadleaf mixed stands and four deciduous broadleaf stands, respectively, in eastern Northeast China. The carbon (C), nitrogen (N), and phosphorus (P) contents of both soil by horizons, litterfall and litter were determined, as well as the soil properties such as pH and bulk density.
Important findings The results showed that soil C, N, and P contents decreased with the soil depth in the RG and TR. The soil C, N and P contents of the O (organic) horizon varied from 53.78 to 90.59 g·kg^-1, 5.02 to 7.83 g·kg^-1, and 0.75 to 0.91 g·kg^-1in all the stands, respectively. The soil C and N contents of the RG were significantly lower than those of the TR in the O horizon, while the soil C, N and P contents of the RG were higher than those of the TR in the A (humus) and B (illuvial) horizons. The soil C density of the O horizon in the RG was significantly lower than that in the TR. The C:N, C:P, and N:P varied from 10.08 to 12.53, 43.97 to 135.52, and 4.56 to 11.64 in all the stands, respectively. The C:N of the O horizon had no significant difference between RG and TR, but the C:P and N:P of the O and A horizons in the RG were significantly lower than those in the TR. There was a significant positive correlation between C and N contents in all soil horizons (R² ranged from 0.40 to 0.76). Except for C:N, restoration approaches, soil horizons and their interaction had significant effects on the content, density and stoichiometry of C, N and P in soil. Soil bulk density and litterfall C content significantly affected soil C, N and P contents. These findings suggest that increasing the proportion of coniferous species by reforestation reduced the content of C and N in the topsoil, which decreased the carbon sequestration accordingly. However, restoration approaches had no significant effect on the C:N, featuring a relatively stable carbon and nitrogen stoichiometry.

Key words: C:N:P, ecological stoichiometry, forest restoration, nutrient cycling, soil carbon sequestration

王燕, 张全智, 王传宽, 郭万桂, 蔺佳玮. 恢复方式对东北东部森林土壤碳氮磷计量特征的影响. 植物生态学报, 2024, 48(7): 943-954. DOI: 10.17521/cjpe.2023.0234

WANG Yan, ZHANG Quan-Zhi, WANG Chuan-Kuan, GUO Wan-Gui, LIN Jia-Wei. Effects of restoration approaches on forest soil carbon, nitrogen and phosphorus stoichiometry in eastern Northeast China. Chinese Journal of Plant Ecology, 2024, 48(7): 943-954. DOI: 10.17521/cjpe.2023.0234

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2023.0234

https://www.plant-ecology.com/CN/Y2024/V48/I7/943

图/表 8

表1 不同恢复方式下东北东部森林类型立地条件及林分特征综述(平均值±标准误, n = 3)

Table 1 Site conditions and stand characteristics of forest types under different restoration approaches in eastern Northeast China (mean ± SE, n = 3)

恢复方式 Restoration approach	林分类型 Forest stand	坡位 Location	坡度 Slope (°)	林龄 Stand age (a)	林分密度 Stand density (trees·hm^-2)	胸高断面积 Basal area at breast height (m²·hm^-2)	平均胸径 Mean DBH (cm)	树种组成 Overstory composition
人工恢复 Artificial restoration	红松林 Pinus koraiensis plantation	中坡位 Mid-slope	12	56	1 989 ± 123	47.34 ± 0.32	16.22 ± 0.32	7PK 3BP
	落叶松林 Larix gmelinii plantation	下坡位 Toe-slope	3	64	739 ± 224	32.93 ± 4.09	20.30 ± 1.28	9LG 1UP
	水曲柳-落叶松混交林 Fraxinus mandshurica - Larix gmelinii mixed plantation	中坡位 Mid-slope	12	57	1 711 ± 58	38.08 ± 2.74	14.00 ± 0.54	6LG 3FM 1UP
天然恢复 Natural restoration	硬阔叶林 Hardwood stand	下坡位 Toe-slope	7	67	1 300 ± 562	35.12 ± 1.58	13.62 ± 0.82	6FM 2JM 1QM 1UP + AM + PA
	杨桦林 Aspen-birch stand	中坡位 Mid-slope	16	72	1 478 ± 42	32.16 ± 3.81	13.47 ± 0.63	5PD 1FM 1TA 1AM 1QM 1BP + JM + PA
	杂木林 Mixed deciduous stand	中坡位 Mid-slope	14	72	1 722 ± 302	35.61 ± 1.45	12.76 ± 0.60	2JM 2PD 1PA 1TA 1AM 1QM 1UP 1BP
	蒙古栎林 Quercus mongolica stand	上坡位 Upper slope	23	73	1 678 ± 82	37.43 ± 1.13	13.83 ± 0.72	10QM + TA + UP

表2 不同恢复方式下东北东部森林土壤的基本性状

Table 2 Basic properties of the forest soil under two restoration approaches in eastern Northeast China

土壤发生层 Soil horizon	恢复方式 Restoration approach	土壤厚度 Soil thickness (cm)	土壤密度 Soil bulk density (g·cm^-3)
O	RG	7.49 ± 0.33	0.73 ± 0.02^***
O	TR	7.83 ± 0.32	0.59 ± 0.02^***
A	RG	20.70 ± 0.58	0.95 ± 0.02
A	TR	21.17 ± 0.43	0.96 ± 0.02
B	RG	25.86 ± 0.96	1.14 ± 0.03^***
B	TR	25.58 ± 0.51	1.31 ± 0.02^***

图1 不同恢复方式东北东部森林类型间土壤碳(C)、氮(N)和磷(P)含量比较。RG, 人工恢复; TR, 天然恢复。A, 腐殖质层; B, 淀积层; O, 有机质层。散点图表示各林型的C、N和P含量在不同土壤深度下的分布情况。不同小写字母表示同一恢复方式森林类型不同土壤发生层间差异显著(p < 0.05 )。**, p < 0.01; ***, p < 0.001; ns, p > 0.05。

Fig. 1 Comparisons of carbon (C), nitrogen (N) and phosphorus (P) contents in forest soil between the two restoration approaches in eastern Northeast China. RG, artificial restoration; TR, natural restoration. A, humus horizon; B, illuvial horizon; O, organic horizon. The scatter plot shows the distribution of C, N and P contents in different soil depths. Different lowercase letters stand for significant difference among the soil horizon of the same restoration approaches (p < 0.05). **, p < 0.01; ***, p < 0.001; ns, p > 0.05.

图2 不同恢复方式东北东部森林类型间土壤各发生层的碳(C)、氮(N)和磷(P)密度的差异性。RG, 人工恢复; TR, 天然恢复。A, 腐殖质层; B, 淀积层; O, 有机质层。不同小写字母表示同一恢复方式森林类型不同土壤发生层间差异显著(p < 0.05)。**, p < 0.01; ***, p < 0.001; ns, p > 0.05。

Fig. 2 Differences in forest soil carbon (C), nitrogen (N) and phosphorus (P) density between the two restoration approaches in eastern Northeast China. RG, artificial restoration; TR, natural restoration. A, humus horizon; B, illuvial horizon; O, organic horizon. Different lowercase letters stand for significant difference among the soil horizon of the same restoration approaches (p < 0.05). **, p < 0.01; ***, p < 0.001; ns, p > 0.05.

图3 不同恢复方式东北东部森林类型间土壤碳氮比、碳磷比和氮磷比差异性。RG, 人工恢复; TR, 天然恢复。A, 腐殖质层; B, 淀积层; O, 有机质层。不同小写字母表示同一恢复方式森林类型不同土壤发生层间差异显著(p < 0.05); *, p < 0.05; **, p < 0.01; ***, p < 0.001; ns, p > 0.05。

Fig. 3 Differences in soil carbon (C):nitrogen (N), C:phosphorus (P) and N:P between the two restoration approaches in eastern Northeast China. RG, artificial restoration; TR, natural restoration. A, humus horizon; B, illuvial horizon; O, organic horizon. Different lowercase letters stand for significant difference among the soil horizon of the same restoration approaches (p < 0.05). *, p < 0.05; **, p < 0.01; ***, p < 0.001; ns, p > 0.05.

图4 不同恢复方式东北东部森林类型间各个土壤发生层的碳(C)-氮(N)、C-磷(P)和N-P含量的相关性分析。RG, 人工恢复; TR, 天然恢复。A-C, O (有机质)层土壤发生层。D-F, A (腐殖质)层土壤发生层。G-I, B (淀积)层土壤发生层。

Fig. 4 Correlation analysis of carbon (C)-nitrogen (N), C-phosphorus (P), and N-P contents by soil horizon between the two restoration approaches in eastern Northeast China. RG, artificial restoration; TR, natural restoration. A-C, soil O (organic) horizon. D-F, soil A (humus) horizon. G-I, soil B (illuvial) horizon.

表3 恢复方式和土壤发生层对东北东部森林土壤碳(C)、氮(N)和磷(P)化学计量影响的方差分析

Table 3 ANOVA of the effects of restoration approaches and soil horizon on forest soil carbon (C), nitrogen (N) and phosphorus (P) stoichiometry in eastern Northeast China

指标 Indicator	处理方式 Treatment	自由度 df	均方误差 MSE	F	p
碳含量 C content	FT	1	977	12.95	***
	SL	2	27 282	316.38	***
	FT × SL	2	3 151	41.74	***
氮含量 N content	FT	1	2.73	6.18	*
	SL	2	197.78	448.03	***
	FT × SL	2	21.94	49.70	***
磷含量 P content	FT	1	0.67	36.25	***
	SL	2	1.75	94.28	***
	FT × SL	2	0.06	3.14	*
碳密度 C density	FT	1	619	5.06	*
	SL	2	3 261	26.69	***
	FT × SL	2	1 385	11.34	***
氮密度 N density	FT	1	12.74	16.67	***
	SL	2	45.31	59.26	***
	FT × SL	2	8.05	10.53	***
磷密度 P density	FT	1	2.74	40.57	***
	SL	2	4.41	65.35	***
	FT × SL	2	0.24	3.58	*
C:N	FT	1	12.48	2.81	ns
	SL	2	5.58	1.26	ns
	FT × SL	2	7.60	1.71	ns
C:P	FT	1	20 436	30.17	***
	SL	2	13 560	20.02	***
	FT × SL	2	7 522	11.10	***
N:P	FT	1	165.66	24.34	***
	SL	2	59.72	8.77	***
	FT × SL	2	27.33	4.02	*

图5 土壤碳(C)、氮(N)和磷(P)含量的影响因子的冗余分析(RDA)。DC, 凋落物C含量; DN, 凋落物N含量; DP, 凋落物P含量; KC, 枯落物C含量; KN, 枯落物N含量; KP, 枯落物P含量; pH, 土壤pH; RZ, 土壤密度; SW, 土壤含水率; TC, 土壤C含量; TN, 土壤N含量; TP, 土壤P含量。

Fig. 5 Redundancy analysis (RDA) of the influencing factors of the soil carbon (C), nitrogen (N) and phosphorus (P) content. DC, litterfall C content; DN, litterfall N content; DP, litterfall P content; KC, litter C content; KN, litter N content; KP, litter P content; pH, soil pH; RZ, soil bulk density; SW, soil moisture; TC, soil C content; TN, soil N content; TP, soil P content.

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