北京东灵山白桦和蒙古栎的根际效应及其季节动态

doi:10.17521/cjpe.2022.0485

摘要/Abstract

摘要：

为了探究木本植物根际效应的季节动态及其驱动因素, 以北京市东灵山地区两种主要植被类型——白桦(Betula platyphylla)林和蒙古栎(Quercus mongolica)林中的优势树种为研究对象, 于2017年的春(5月)、夏(7月)、秋(9月)、冬(12月) 4个季节分别测定根际土壤与非根际土壤的理化性质、微生物生物量、碳矿化速率及净氮矿化速率、胞外酶活性和矢量特征以及植物的根系、叶片功能性状, 分析根际效应的季节变化动态。结果发现: (1)土壤pH及铵态氮含量、微生物生物量、碳氮矿化速率、胞外酶活性和矢量特征指标在根际土壤与非根际土壤之间存在显著差异, 根际效应主要呈现为正效应, 即根际土壤高于非根际土壤。(2)根际效应存在显著的季节动态, 表现为秋季的根际效应最强。(3)根际效应与植物根系和叶片功能性状之间存在显著相关关系。细根生物量与可提取有机碳、土壤总碳、总氮含量的根际效应显著正相关; 叶干物质含量、叶碳氮比与微生物生物量碳含量、微生物生物量氮含量、碳矿化速率、酸性磷酸酶活性的根际效应显著正相关。研究结果表明, 植物功能性状对于植物的根际效应具有重要作用; 在东灵山的温带落叶阔叶林, 可能由于秋季植物地下碳输入量最高, 导致根际微生物数量和活性增加, 从而出现秋季微生物生物量及活性的根际效应高于其他季节的现象。

关键词: 根际效应, 季节动态, 植物功能性状, 土壤胞外酶, 森林

Abstract:

Aims The objective of this study was to explore the seasonal variations of the rhizosphere effects of woody plants and their driving factors, and to assess the importance of plant functional traits in the control of rhizosphere processes.
Methods We collected paired rhizosphere and bulk soils of the dominant tree species of two main types of vegetation in Dongling Mountain, Beijing, Betula platyphylla forest and Quercus mongolica forest. Soil physiochemical properties, microbial biomass, carbon and net nitrogen mineralization rates, extracellular enzyme activities and vector characteristics of rhizosphere and bulk soils, as well as plant root and leaf functional traits, in spring (May), summer (July), autumn (September), and winter (December) of 2017 were measured to analyze the seasonal dynamics of rhizosphere effects and their driving factors.
Important findings (1) There were significant differences in soil pH, NH₄⁺-N, microbial biomass, carbon and net nitrogen mineralization rates, extracellular enzyme activities and vector characteristics between rhizosphere soil and bulk soil, and these rhizosphere effects were mainly positive. (2) The rhizosphere effects had significant seasonal dynamics, usually being strongest in autumn. (3) There were often significant correlations between rhizosphere effects and plant root and leaf functional traits. Among them, fine root biomass was significantly and positively correlated with the rhizosphere effect on contents of extractable organic carbon, soil total carbon and total nitrogen. Leaf dry matter content and leaf carbon and nitrogen ratio were significantly and positively correlated with the rhizosphere effect on microbial biomass carbon content, microbial biomass nitrogen content, carbon mineralization rate, and acid phosphatase activity. These results showed that the functional traits of plants were of great significance in rhizosphere processes. In the temperate deciduous broadleaf forest in Dongling Mountain, the highest belowground carbon allocation of plants leads to an increase in the biomass and activity of rhizosphere microorganisms in autumn, which makes the rhizosphere effect of microbial biomass and activity in autumn higher than that in other seasons.

Key words: rhizosphere effect, seasonal dynamic, plant functional traits, soil extracellular enzymes, forest

付粱晨, 丁宗巨, 唐茂, 曾辉, 朱彪. 北京东灵山白桦和蒙古栎的根际效应及其季节动态. 植物生态学报, 2024, 48(4): 508-522. DOI: 10.17521/cjpe.2022.0485

FU Liang-Chen, DING Zong-Ju, TANG Mao, ZENG Hui, ZHU Biao. Rhizosphere effects of Betula platyphylla and Quercus mongolica and their seasonal dynamics in Dongling Mountain, Beijing. Chinese Journal of Plant Ecology, 2024, 48(4): 508-522. DOI: 10.17521/cjpe.2022.0485

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2022.0485

https://www.plant-ecology.com/CN/Y2024/V48/I4/508

图/表 7

表1 北京东灵山白桦林和蒙古栎林土壤理化性质、微生物生物量、碳氮矿化速率、胞外酶活性及矢量特征基本信息(平均值±标准误, n = 4)

Table 1 Soil physiochemical properties, microbial biomass, carbon and nitrogen mineralization rates, soil extracellular enzyme activities and vector characteristics of Betula platyphylla forest and Quercus mongolica forest in Dongling Mountain, Beijing (mean ± SE, n = 4)

		白桦林 Betula platyphylla forest					蒙古栎林 Quercus mongolica forest
		春 Spring	夏 Summer	秋 Autumn		冬 Winter	春 Spring		夏 Summer	秋 Autumn	冬 Winter
SWC (%)	r	42.11 ± 2.97^a	45.71 ± 1.39^a	18.77 ± 1.04^b		45.74 ± 5.75^a	17.11 ± 0.39^c		30.75 ± 1.18^a	13.32 ± 0.80^d	26.13 ± 1.46^b
SWC (%)	b	43.62 ± 2.20^a	44.85 ± 1.21^a	16.84 ± 0.76^b		46.25 ± 7.61^a	20.96 ± 3.26^b		29.76 ± 1.62^a	12.60 ± 0.84^c	24.33 ± 1.85a^b
pH	r	5.48 ± 0.08^c	5.97 ± 0.19^b	6.45 ± 0.16^a		5.89 ± 0.07^bc	5.64 ± 0.07^b		5.55 ± 0.13^b	6.20 ± 0.17^a	5.57 ± 0.19^b
pH	b	6.41 ± 0.06^a	6.39 ± 0.24^a	6.65 ± 0.14^a		6.46 ± 0.13^a	6.34 ± 0.08^a		5.82 ± 0.06^b	6.44 ± 0.13^a	5.55 ± 0.21^b
EOC content (mg·kg^-1)	r	1 420.02 ± 212.81^a	1 552.22 ± 143.35^a	741.49 ± 125.89^b		491.32 ± 110.09^b	955.42 ± 77.67^a		1 092.27 ± 81.65^a	494.17 ± 43.51^b	263.94 ± 20.23^c
EOC content (mg·kg^-1)	b	1 175.09 ± 138.55^a	1 289.81 ± 96.76^a	485.55 ± 69.81^b		437.68 ± 74.89^b	922.87 ± 113.96^a		1 092.05 ± 102.71^a	404.52 ± 45.92^b	242.67 ± 13.41^b
TC content (mg·g^-1)	r	65.97 ± 6.84^a	63.22 ± 3.51^a	58.08 ± 5.95^a		88.08 ± 20.58^a	37.54 ± 1.03^a		35.58 ± 1.36^a	39.83 ± 3.33^a	35.97 ± 2.61^a
TC content (mg·g^-1)	b	58.99 ± 5.69^ab	55.09 ± 2.39^b	46.01 ± 4.06^b		86.64 ± 18.54^a	35.73 ± 1.74^a		33.94 ± 2.37^a	38.47 ± 3.58^a	34.64 ± 2.60^a
TN content (mg·g^-1)	r	5.16 ± 0.43^a	5.19 ± 0.32^a	4.54 ± 0.43^a		6.65 ± 1.49^a	3.18 ± 0.06^a		2.95 ± 0.14^a	3.27 ± 0.24^a	3.01 ± 0.23^a
TN content (mg·g^-1)	b	4.73 ± 0.42^ab	4.55 ± 0.23^ab	3.79 ± 0.27^b		6.65 ± 1.38^a	3.12 ± 0.19^a		2.85 ± 0.23^a	3.15 ± 0.27^a	2.91 ± 0.22^a
Soil C:N	r	12.74 ± 0.35^ab	12.19 ± 0.15^b	12.76 ± 0.13^ab		13.10 ± 0.35^a	11.81 ± 0.32^a		12.07 ± 0.30^a	12.16 ± 0.23^a	11.96 ± 0.13^a
Soil C:N	b	12.45 ± 0.13^b	12.11 ± 0.10^b	12.10 ± 0.21^b		12.96 ± 0.13^a	11.48 ± 0.30^a		11.97 ± 0.31^a	12.19 ± 0.16^a	11.91 ± 0.08^a
ETN content (mg·kg^-1)	r	138.18 ± 16.83^a	135.56 ± 10.73^a	50.48 ± 11.13^b		67.00 ± 15.17^b	86.18 ± 5.38^b		100.25 ± 2.12^a	31.80 ± 2.50^c	31.89 ± 1.82^c
ETN content (mg·kg^-1)	b	100.97 ± 11.66^a	110.43 ± 5.31^a	39.88 ± 5.38^b		59.03 ± 8.36^b	82.00 ± 8.12^a		93.73 ± 4.49^a	28.51 ± 1.57^b	33.99 ± 0.88^b
NH₄⁺-N content (μg·g^-1)	r	15.14 ± 1.34^ab	11.63 ± 1.15^ab	9.55 ± 1.24^b		19.79 ± 4.99^a	11.43 ± 0.54^ab		15.41 ± 2.54^a	8.53 ± 0.58^b	7.52 ± 0.42^b
NH₄⁺-N content (μg·g^-1)	b	5.08 ± 0.36^b	7.75 ± 0.68^a	6.86 ± 1.01^ab		7.18 ± 0.72^ab	6.10 ± 0.97^b		10.12 ± 1.34^a	5.02 ± 0.12^b	6.29 ± 0.71^b
NO₃^--N content (μg·g^-1)	r	2.94 ± 0.32^bc	5.82 ± 1.46^b	0.12 ± 0.02^c		12.43 ± 2.72^a	1.31 ± 0.28^b		4.57 ± 0.74^a	0.07 ± 0.00^b	3.98 ± 0.31^a
NO₃^--N content (μg·g^-1)	b	5.99 ± 0.76^b	7.31 ± 0.87^b	0.00 ± 0.00^c		16.42 ± 3.43^a	2.60 ± 0.67^b		5.99 ± 0.85^a	0.00 ± 0.00^c	5.14 ± 1.15^a
MBC content (mg·kg^-1)	r	1 224.41 ± 71.60^a	1 456.06 ± 115.29^a	1 202.39 ± 118.77^a		1 362.10 ± 290.82^a	509.07 ± 60.41^c		732.73 ± 39.53^b	920.16 ± 73.95^a	447.73 ± 40.34^c
MBC content (mg·kg^-1)	b	1 222.59 ± 140.84^a	1 160.56 ± 72.54^ab	692.14 ± 66.88^b		1 048.46 ± 295.22^ab	648.17 ± 104.11^a		608.29 ± 54.09^ab	405.67 ± 46.44^b	487.32 ± 71.78^ab
MBN content (mg·kg^-1)	r	210.49 ± 5.73^a	225.58 ± 19.78^a	86.91 ± 11.09^b		137.24 ± 29.16^b	66.34 ± 5.34^b		105.61 ± 7.91^a	62.91 ± 9.23^b	40.69 ± 4.47^c
MBN content (mg·kg^-1)	b	187.31 ± 26.82^a	187.49 ± 19.21^a	67.81 ± 7.86^b		140.38 ± 41.66^ab	93.36 ± 14.70^a		100.08 ± 6.35^a	36.12 ± 0.86^b	51.42 ± 9.80^b
MBC:MBN	r	5.82 ± 0.31^c	6.48 ± 0.24^c	14.03 ± 0.90^a		9.90 ± 0.07^b	7.77 ± 0.91^c		7.00 ± 0.35^c	15.12 ± 1.16^a	11.10 ± 0.62^b
MBC:MBN	b	6.66 ± 0.41^b	6.27 ± 0.29^b	10.38 ± 0.87^a		7.95 ± 0.72^b	7.03 ± 0.74^b		6.07 ± 0.38^b	11.18 ± 1.12^a	9.80 ± 0.68^a
C_min(mg·g^-1·d^-1)	r	0.21 ± 0.02^a	0.17 ± 0.01^a	0.04 ± 0.01^c		0.10 ± 0.01^b	0.14 ± 0.01^a		0.12 ± 0.02^a	0.02 ± 0.00^b	0.04 ± 0.00^b
C_min(mg·g^-1·d^-1)	b	0.21 ± 0.02^a	0.16 ± 0.02^b	0.02 ± 0.00^c		0.05 ± 0.01^c	0.18 ± 0.01^a		0.11 ± 0.01^b	0.02 ± 0.00^c	0.03 ± 0.00^c
N_min(μg·g^-1·d^-1)	r	5.01 ± 0.72^a	1.89 ± 0.29^b	0.98 ± 0.21^b		1.62 ± 0.35^b	1.02 ± 0.30^a		0.85 ± 0.23^ab	0.33 ± 0.08^b	0.90 ± 0.12^ab
N_min(μg·g^-1·d^-1)	b	3.23 ± 0.42^a	1.85 ± 0.26^b	0.81 ± 0.11^c		1.68 ± 0.23^b	1.38 ± 0.26^a		1.02 ± 0.37^ab	0.33 ± 0.06^b	1.07 ± 0.08^ab
	r	6.80 ± 1.58^b	21.89 ± 5.09^a	14.84 ± 1.97^ab		8.17 ± 1.61^b	2.68 ± 0.73^b		8.42 ± 0.59^a	6.48 ± 0.81^a	2.07 ± 0.43^b
BG activity (nmol·g^-1·h^-1)	b	13.34 ± 2.19^b	26.06 ± 4.15^a	16.24 ± 2.72^b		12.19 ± 1.84^b	4.95 ± 0.80^c		12.94 ± 1.27^a	8.07 ± 0.93^b	3.26 ± 0.53^c
NAG activity (nmol·g^-1·h^-1)	r	5.28 ± 0.60^c	24.51 ± 3.21^a	17.09 ± 1.63^b		6.65 ± 1.63^c	4.49 ± 0.68^b		11.79 ± 0.57^a	10.17 ± 2.57^a	2.74 ± 0.25^b
NAG activity (nmol·g^-1·h^-1)	b	6.62 ± 0.73^c	22.55 ± 1.14^a	14.37 ± 2.54^b		6.02 ± 1.07^c	4.73 ± 0.70^b		10.59 ± 1.03^a	8.68 ± 1.76^a	2.95 ± 0.40^b
AP activity (nmol·g^-1·h^-1)	r	26.79 ± 0.60^b	56.05 ± 5.74^a	36.61 ± 3.30^b		26.93 ± 3.61^b	19.43 ± 1.11^c		48.05 ± 1.25^a	32.79 ± 3.03^b	20.54 ± 1.60^c
AP activity (nmol·g^-1·h^-1)	b	29.90 ± 1.20^b	57.40 ± 11.50^a	32.49 ± 2.06^b		25.01 ± 1.76^b	22.56 ± 1.91^b		43.32 ± 4.59^a	25.08 ± 1.78^b	25.63 ± 1.42^b
POX+PER activity (μmol·g^-1·h^-1)	r	22.87 ± 1.69^a	10.10 ± 2.03^b	8.55 ± 1.06^b	11.23 ± 1.26^b		21.98 ± 3.58^a	14.08 ± 0.74^b		12.22 ± 1.58^b	13.07 ± 1.05^b
POX+PER activity (μmol·g^-1·h^-1)	b	24.52 ± 0.67^a	10.50 ± 1.54^b	8.23 ± 1.66^b	11.79 ± 1.08^b		21.06 ± 4.10^a	12.97 ± 0.76^bc		6.14 ± 0.92^c	15.83 ± 2.96^ab
Vector length	r	0.58 ± 0.05^a	0.54 ± 0.06^a	0.55 ± 0.03^a	0.61 ± 0.01^a		0.38 ± 0.06^a	0.44 ± 0.03^a		0.44 ± 0.05^a	0.43 ± 0.04^a
Vector length	b	0.73 ± 0.03^a	0.63 ± 0.06^a	0.62 ± 0.05^a	0.75 ± 0.01^a		0.54 ± 0.03^a	0.60 ± 0.01^a		0.55 ± 0.01^a	0.53 ± 0.06^a
Vector angle	r	70.62 ± 2.37^a	59.84 ± 3.90^bc	58.25 ± 2.46^c	67.96 ± 1.82^ab		72.20 ± 2.91^ab	70.31 ± 0.30^b		67.60 ± 1.87^b	77.84 ± 1.83^a
Vector angle	b	65.54 ± 1.54^a	59.08 ± 4.07^a	58.22 ± 1.98^a	64.48 ± 1.98^a		70.67 ± 2.47^b	67.21 ± 1.37^bc		63.50 ± 2.85^c	77.77 ± 1.45^a

表1 北京东灵山白桦林和蒙古栎林土壤理化性质、微生物生物量、碳氮矿化速率、胞外酶活性及矢量特征基本信息(平均值±标准误, n = 4)

		白桦林 Betula platyphylla forest					蒙古栎林 Quercus mongolica forest
		春 Spring	夏 Summer	秋 Autumn		冬 Winter	春 Spring		夏 Summer	秋 Autumn	冬 Winter
SWC (%)	r	42.11 ± 2.97^a	45.71 ± 1.39^a	18.77 ± 1.04^b		45.74 ± 5.75^a	17.11 ± 0.39^c		30.75 ± 1.18^a	13.32 ± 0.80^d	26.13 ± 1.46^b
SWC (%)	b	43.62 ± 2.20^a	44.85 ± 1.21^a	16.84 ± 0.76^b		46.25 ± 7.61^a	20.96 ± 3.26^b		29.76 ± 1.62^a	12.60 ± 0.84^c	24.33 ± 1.85a^b
pH	r	5.48 ± 0.08^c	5.97 ± 0.19^b	6.45 ± 0.16^a		5.89 ± 0.07^bc	5.64 ± 0.07^b		5.55 ± 0.13^b	6.20 ± 0.17^a	5.57 ± 0.19^b
pH	b	6.41 ± 0.06^a	6.39 ± 0.24^a	6.65 ± 0.14^a		6.46 ± 0.13^a	6.34 ± 0.08^a		5.82 ± 0.06^b	6.44 ± 0.13^a	5.55 ± 0.21^b
EOC content (mg·kg^-1)	r	1 420.02 ± 212.81^a	1 552.22 ± 143.35^a	741.49 ± 125.89^b		491.32 ± 110.09^b	955.42 ± 77.67^a		1 092.27 ± 81.65^a	494.17 ± 43.51^b	263.94 ± 20.23^c
EOC content (mg·kg^-1)	b	1 175.09 ± 138.55^a	1 289.81 ± 96.76^a	485.55 ± 69.81^b		437.68 ± 74.89^b	922.87 ± 113.96^a		1 092.05 ± 102.71^a	404.52 ± 45.92^b	242.67 ± 13.41^b
TC content (mg·g^-1)	r	65.97 ± 6.84^a	63.22 ± 3.51^a	58.08 ± 5.95^a		88.08 ± 20.58^a	37.54 ± 1.03^a		35.58 ± 1.36^a	39.83 ± 3.33^a	35.97 ± 2.61^a
TC content (mg·g^-1)	b	58.99 ± 5.69^ab	55.09 ± 2.39^b	46.01 ± 4.06^b		86.64 ± 18.54^a	35.73 ± 1.74^a		33.94 ± 2.37^a	38.47 ± 3.58^a	34.64 ± 2.60^a
TN content (mg·g^-1)	r	5.16 ± 0.43^a	5.19 ± 0.32^a	4.54 ± 0.43^a		6.65 ± 1.49^a	3.18 ± 0.06^a		2.95 ± 0.14^a	3.27 ± 0.24^a	3.01 ± 0.23^a
TN content (mg·g^-1)	b	4.73 ± 0.42^ab	4.55 ± 0.23^ab	3.79 ± 0.27^b		6.65 ± 1.38^a	3.12 ± 0.19^a		2.85 ± 0.23^a	3.15 ± 0.27^a	2.91 ± 0.22^a
Soil C:N	r	12.74 ± 0.35^ab	12.19 ± 0.15^b	12.76 ± 0.13^ab		13.10 ± 0.35^a	11.81 ± 0.32^a		12.07 ± 0.30^a	12.16 ± 0.23^a	11.96 ± 0.13^a
Soil C:N	b	12.45 ± 0.13^b	12.11 ± 0.10^b	12.10 ± 0.21^b		12.96 ± 0.13^a	11.48 ± 0.30^a		11.97 ± 0.31^a	12.19 ± 0.16^a	11.91 ± 0.08^a
ETN content (mg·kg^-1)	r	138.18 ± 16.83^a	135.56 ± 10.73^a	50.48 ± 11.13^b		67.00 ± 15.17^b	86.18 ± 5.38^b		100.25 ± 2.12^a	31.80 ± 2.50^c	31.89 ± 1.82^c
ETN content (mg·kg^-1)	b	100.97 ± 11.66^a	110.43 ± 5.31^a	39.88 ± 5.38^b		59.03 ± 8.36^b	82.00 ± 8.12^a		93.73 ± 4.49^a	28.51 ± 1.57^b	33.99 ± 0.88^b
NH₄⁺-N content (μg·g^-1)	r	15.14 ± 1.34^ab	11.63 ± 1.15^ab	9.55 ± 1.24^b		19.79 ± 4.99^a	11.43 ± 0.54^ab		15.41 ± 2.54^a	8.53 ± 0.58^b	7.52 ± 0.42^b
NH₄⁺-N content (μg·g^-1)	b	5.08 ± 0.36^b	7.75 ± 0.68^a	6.86 ± 1.01^ab		7.18 ± 0.72^ab	6.10 ± 0.97^b		10.12 ± 1.34^a	5.02 ± 0.12^b	6.29 ± 0.71^b
NO₃^--N content (μg·g^-1)	r	2.94 ± 0.32^bc	5.82 ± 1.46^b	0.12 ± 0.02^c		12.43 ± 2.72^a	1.31 ± 0.28^b		4.57 ± 0.74^a	0.07 ± 0.00^b	3.98 ± 0.31^a
NO₃^--N content (μg·g^-1)	b	5.99 ± 0.76^b	7.31 ± 0.87^b	0.00 ± 0.00^c		16.42 ± 3.43^a	2.60 ± 0.67^b		5.99 ± 0.85^a	0.00 ± 0.00^c	5.14 ± 1.15^a
MBC content (mg·kg^-1)	r	1 224.41 ± 71.60^a	1 456.06 ± 115.29^a	1 202.39 ± 118.77^a		1 362.10 ± 290.82^a	509.07 ± 60.41^c		732.73 ± 39.53^b	920.16 ± 73.95^a	447.73 ± 40.34^c
MBC content (mg·kg^-1)	b	1 222.59 ± 140.84^a	1 160.56 ± 72.54^ab	692.14 ± 66.88^b		1 048.46 ± 295.22^ab	648.17 ± 104.11^a		608.29 ± 54.09^ab	405.67 ± 46.44^b	487.32 ± 71.78^ab
MBN content (mg·kg^-1)	r	210.49 ± 5.73^a	225.58 ± 19.78^a	86.91 ± 11.09^b		137.24 ± 29.16^b	66.34 ± 5.34^b		105.61 ± 7.91^a	62.91 ± 9.23^b	40.69 ± 4.47^c
MBN content (mg·kg^-1)	b	187.31 ± 26.82^a	187.49 ± 19.21^a	67.81 ± 7.86^b		140.38 ± 41.66^ab	93.36 ± 14.70^a		100.08 ± 6.35^a	36.12 ± 0.86^b	51.42 ± 9.80^b
MBC:MBN	r	5.82 ± 0.31^c	6.48 ± 0.24^c	14.03 ± 0.90^a		9.90 ± 0.07^b	7.77 ± 0.91^c		7.00 ± 0.35^c	15.12 ± 1.16^a	11.10 ± 0.62^b
MBC:MBN	b	6.66 ± 0.41^b	6.27 ± 0.29^b	10.38 ± 0.87^a		7.95 ± 0.72^b	7.03 ± 0.74^b		6.07 ± 0.38^b	11.18 ± 1.12^a	9.80 ± 0.68^a
C_min(mg·g^-1·d^-1)	r	0.21 ± 0.02^a	0.17 ± 0.01^a	0.04 ± 0.01^c		0.10 ± 0.01^b	0.14 ± 0.01^a		0.12 ± 0.02^a	0.02 ± 0.00^b	0.04 ± 0.00^b
C_min(mg·g^-1·d^-1)	b	0.21 ± 0.02^a	0.16 ± 0.02^b	0.02 ± 0.00^c		0.05 ± 0.01^c	0.18 ± 0.01^a		0.11 ± 0.01^b	0.02 ± 0.00^c	0.03 ± 0.00^c
N_min(μg·g^-1·d^-1)	r	5.01 ± 0.72^a	1.89 ± 0.29^b	0.98 ± 0.21^b		1.62 ± 0.35^b	1.02 ± 0.30^a		0.85 ± 0.23^ab	0.33 ± 0.08^b	0.90 ± 0.12^ab
N_min(μg·g^-1·d^-1)	b	3.23 ± 0.42^a	1.85 ± 0.26^b	0.81 ± 0.11^c		1.68 ± 0.23^b	1.38 ± 0.26^a		1.02 ± 0.37^ab	0.33 ± 0.06^b	1.07 ± 0.08^ab
	r	6.80 ± 1.58^b	21.89 ± 5.09^a	14.84 ± 1.97^ab		8.17 ± 1.61^b	2.68 ± 0.73^b		8.42 ± 0.59^a	6.48 ± 0.81^a	2.07 ± 0.43^b
BG activity (nmol·g^-1·h^-1)	b	13.34 ± 2.19^b	26.06 ± 4.15^a	16.24 ± 2.72^b		12.19 ± 1.84^b	4.95 ± 0.80^c		12.94 ± 1.27^a	8.07 ± 0.93^b	3.26 ± 0.53^c
NAG activity (nmol·g^-1·h^-1)	r	5.28 ± 0.60^c	24.51 ± 3.21^a	17.09 ± 1.63^b		6.65 ± 1.63^c	4.49 ± 0.68^b		11.79 ± 0.57^a	10.17 ± 2.57^a	2.74 ± 0.25^b
NAG activity (nmol·g^-1·h^-1)	b	6.62 ± 0.73^c	22.55 ± 1.14^a	14.37 ± 2.54^b		6.02 ± 1.07^c	4.73 ± 0.70^b		10.59 ± 1.03^a	8.68 ± 1.76^a	2.95 ± 0.40^b
AP activity (nmol·g^-1·h^-1)	r	26.79 ± 0.60^b	56.05 ± 5.74^a	36.61 ± 3.30^b		26.93 ± 3.61^b	19.43 ± 1.11^c		48.05 ± 1.25^a	32.79 ± 3.03^b	20.54 ± 1.60^c
AP activity (nmol·g^-1·h^-1)	b	29.90 ± 1.20^b	57.40 ± 11.50^a	32.49 ± 2.06^b		25.01 ± 1.76^b	22.56 ± 1.91^b		43.32 ± 4.59^a	25.08 ± 1.78^b	25.63 ± 1.42^b
POX+PER activity (μmol·g^-1·h^-1)	r	22.87 ± 1.69^a	10.10 ± 2.03^b	8.55 ± 1.06^b	11.23 ± 1.26^b		21.98 ± 3.58^a	14.08 ± 0.74^b		12.22 ± 1.58^b	13.07 ± 1.05^b
POX+PER activity (μmol·g^-1·h^-1)	b	24.52 ± 0.67^a	10.50 ± 1.54^b	8.23 ± 1.66^b	11.79 ± 1.08^b		21.06 ± 4.10^a	12.97 ± 0.76^bc		6.14 ± 0.92^c	15.83 ± 2.96^ab
Vector length	r	0.58 ± 0.05^a	0.54 ± 0.06^a	0.55 ± 0.03^a	0.61 ± 0.01^a		0.38 ± 0.06^a	0.44 ± 0.03^a		0.44 ± 0.05^a	0.43 ± 0.04^a
Vector length	b	0.73 ± 0.03^a	0.63 ± 0.06^a	0.62 ± 0.05^a	0.75 ± 0.01^a		0.54 ± 0.03^a	0.60 ± 0.01^a		0.55 ± 0.01^a	0.53 ± 0.06^a
Vector angle	r	70.62 ± 2.37^a	59.84 ± 3.90^bc	58.25 ± 2.46^c	67.96 ± 1.82^ab		72.20 ± 2.91^ab	70.31 ± 0.30^b		67.60 ± 1.87^b	77.84 ± 1.83^a
Vector angle	b	65.54 ± 1.54^a	59.08 ± 4.07^a	58.22 ± 1.98^a	64.48 ± 1.98^a		70.67 ± 2.47^b	67.21 ± 1.37^bc		63.50 ± 2.85^c	77.77 ± 1.45^a

表2 北京东灵山白桦和蒙古栎植物根叶功能性状基本信息(平均值±标准误, n = 4)

Table 2 Plant functional traits of leaves and roots of Betula platyphylla and Quercus mongolica in Dongling Mountain, Beijing (mean ± SE, n = 4)

	白桦 Betula platyphylla				蒙古栎 Quercus mongolica
	春 Spring	夏 Summer	秋 Autumn	冬 Winter	春 Spring	夏 Summer	秋 Autumn	冬 Winter
SLA (m²·kg^-1)	33.1 ± 1.6^a	29.1 ± 2.1^a	22.5 ± 0.8^b	NA	20.4 ± 1.3^a	16.4 ± 0.8^b	16.1 ± 0.3^b	NA
LDMC (g·g^-1)	0.189 ± 0.005^b	0.173 ± 0.008^b	0.261 ± 0.010^a	NA	0.217 ± 0.010^b	0.214 ± 0.004^b	0.348 ± 0.004^a	NA
Leaf C content (mg·g^-1)	NA	481 ± 7^a	491 ± 2^a	NA	458 ± 1^c	480 ± 3^a	467 ± 2^b	NA
Leaf N content (mg·g^-1)	NA	27.4 ± 1.0^a	21.2 ± 0.7^b	NA	56.1 ± 0.7^a	27.5 ± 0.6^b	21.2 ± 0.6^c	NA
Leaf C:N	NA	17.7 ± 0.8^b	23.2 ± 0.8^a	NA	8.2 ± 0.1^c	17.5 ± 0.3^b	22.1 ± 0.5^a	NA
Fine root biomass (g)	21.2 ± 1.8^b	25.6 ± 2.9^b	37.6 ± 2.7^a	6.4 ± 0.3^c	15.9 ± 1.3^b	25.9 ± 1.3^a	28.7 ± 2.8^a	6.7 ± 0.9^c
Root C content (mg·g^-1)	489 ± 8^b	463 ± 7^c	484 ± 6^b	518 ± 3^a	500 ± 4^a	494 ± 8^a	497 ± 9^a	507 ± 1^a
Root N content (mg·g^-1)	12.28 ± 0.47^a	9.98 ± 0.66^ab	9.34 ± 0.32^b	11.19 ± 1.10^ab	9.65 ± 0.61^a	9.13 ± 0.34^a	9.70 ± 0.97^a	8.38 ± 0.69^a
Root C:N	40.1 ± 2.2^b	47.2 ± 3.1^ab	52.1 ± 2.2^a	48.2 ± 4.8^ab	52.7 ± 3.5^a	54.5 ± 2.7^a	53.7 ± 6.2^a	61.9 ± 4.5^a
Fine root density (kg·m^-3)	1 077 ± 127^b	1 285 ± 161^b	2 091 ± 150^a	NA	580 ± 78^b	1437 ± 71^a	1 593 ± 153^a	NA

图1 北京东灵山白桦和蒙古栎林土壤理化性质的根际效应(平均值±标准误, n = 4)的季节变化。A, 土壤含水量。B, pH。C, 可提取有机碳含量。D, 总碳含量。E, 总氮含量。F, 土壤碳氮比。G, 可提取氮含量。H, 铵态氮含量。I, 硝态氮含量。不同小写字母表示同一树种的根际效应在不同季节间差异显著(p < 0.05)。*表示根际效应值显著区别于1 (*, p < 0.05; **, p < 0.01; ***, p < 0.001)。左下角为线性混合效应模型的结果, *表示树种、季节或两者的交互作用对根际效应有显著影响(*, p < 0.05; **, p < 0.01; ***, p < 0.001)。

Fig. 1 Seasonal variations of rhizosphere effects on soil physical and chemical properties of Betula platyphylla forest and Quercus mongolica forest in Dongling Mountain, Beijing (mean ± SE, n = 4). A, Soil water content. B, pH. C, Extractable organic carbon content. D, Total carbon content. E, Total nitrogen content. F, Soil carbon to nitrogen ratio. G, Extractable nitrogen content. H, Ammonium nitrogen content. I, Nitrate nitrogen content. Different lowercase letters indicate the rhizosphere effect of the same species is significantly different in different seasons (p < 0.05). The asterisk indicates that the rhizosphere effect value is significantly different from 1 (*, p < 0.05; **, p < 0.01; ***, p < 0.001). The results of linear mixed-effect model are located in the lower left corner of each figure. The asterisk indicates that tree species, season or their interaction has a significant impact on the rhizosphere effect (*, p < 0.05; **, p < 0.01; ***, p < 0.001).

图2 北京东灵山白桦和蒙古栎林土壤微生物生物量及碳氮矿化速率根际效应(平均值±标准误, n = 4)的季节变化。A, 微生物生物量碳含量。B, 微生物生物量氮含量。C, 微生物生物量碳氮比。D, 碳矿化速率。E, 氮矿化速率。不同小写字母表示同一树种的根际效应在不同季节间差异显著(p < 0.05)。*表示根际效应值显著区别于1 (*, p < 0.05; **, p < 0.01; ***, p < 0.001)。左下角为线性混合效应模型的结果, *表示树种、季节或两者的交互作用对根际效应有显著影响(*, p < 0.05; **, p < 0.01; ***, p < 0.001)。

Fig. 2 Seasonal variations of rhizosphere effects on microbial biomass, carbon and nitrogen mineralization rates of Betula platyphylla forest and Quercus mongolica forest in Dongling Mountain, Beijing (mean ± SE, n = 4). A, Microbial biomass carbon content. B, Microbial biomass nitrogen content. C, Microbial biomass carbon to nitrogen ratio. D, Carbon mineralization rate. E, Nitrogen mineralization rate. Different lowercase letters indicate the rhizosphere effect of the same species is significantly different in different seasons (p < 0.05). The asterisk indicates that the rhizosphere effect value is significantly different from 1 (*, p < 0.05; **, p < 0.01; ***, p < 0.001). The results of linear mixed-effect model are located in the lower left corner of each figure. The asterisk indicates that tree species, season or their interaction has a significant impact on the rhizosphere effect (*, p < 0.05; **, p < 0.01; ***, p < 0.001).

图3 北京东灵山白桦和蒙古栎林土壤胞外酶活性及矢量特征根际效应(平均值±标准误, n = 4)的季节变化。A, β-1,4-葡萄糖苷酶活性。B, β-1,4-乙酰氨基葡糖苷酶活性。C, 酸性磷酸酶活性。D, 氧化酶活性。E, 矢量长度。F, 矢量角度。不同小写字母表示同一树种的根际效应在不同季节间差异显著(p < 0.05)。*表示根际效应值显著区别于1 (*, p < 0.05; **, p < 0.01; ***, p < 0.001)。左下角为线性混合效应模型的结果, *表示树种、季节或两者的交互作用对根际效应有显著影响(*, p < 0.05; **, p < 0.01; ***, p < 0.001)。

Fig. 3 Seasonal variations of rhizosphere effects on soil extracellular enzyme activities and vector characteristics of Betula platyphylla forest and Quercus mongolica forest in Dongling Mountain, Beijing (mean ± SE, n = 4). A, β-1,4-glucosidase. B, β-1,4-acetylglucosidase. C, Acid phosphatase. D, Oxidase activity. E, Vector length. F, Vector angle. Different lowercase letters indicate the rhizosphere effect of the same species is significantly different in different seasons (p < 0.05). The asterisk indicates that the rhizosphere effect value is significantly different from 1 (*, p < 0.05; **, p < 0.01; ***, p < 0.001). The results of linear mixed-effect model are located in the lower left corner of each figure. The asterisk indicates that tree species, season or their interaction has a significant impact on the rhizosphere effect (*, p < 0.05; **, p < 0.01; ***, p < 0.001).

图4 北京东灵山白桦和蒙古栎林根际效应与植物功能性状的相关性分析。A, 白桦。B, 蒙古栎。AP, 酸性磷酸酶活性; BG, β-1,4-葡萄糖苷酶活性; Cmin, 碳矿化速率; ETN, 可提取氮含量; EOC, 可提取有机碳含量; Fine root biomass, 细根生物量; Fine root density, 细根密度; LDMC, 叶干物质含量; Leaf C, 叶碳含量; Leaf N, 叶氮含量; Leaf C:N, 叶碳氮比; MBC, 微生物生物量碳含量; MBN, 微生物生物量氮含量; NAG, β-1,4-乙酰氨基葡糖苷酶活性; NH4+-N, 铵态氮含量; Nmin, 氮矿化速率; NO3--N, 硝态氮含量; POX+PER, 氧化酶活性; Root C, 根碳含量; Root N, 根氮含量; Root C:N, 根碳氮比; SLA, 比叶面积; Soil C:N, 土壤碳氮比; SWC, 土壤含水量; TC, 总碳含量; TN, 总氮含量; Vector length, 矢量长度; Vector angle, 矢量角度。红色代表正相关关系, 蓝色代表负相关关系, 颜色越深, 相关性越强。*表示相关性显著(*, p < 0.05; **, p < 0.01; ***, p < 0.001)。

Fig. 4 Correlation analysis between rhizosphere effect and plant functional traits of Betula platyphylla forest and Quercus mongolica forest in Dongling Mountain, Beijing. A, Betula platyphylla. B, Quercus mongolica. AP, acid phosphatase activity; BG, β-1,4-glucosidase activity; Cmin, carbon mineralization rate; EOC, extractable organic carbon content; ETN, extractable nitrogen content; LDMC, leaf dry matter content; MBC, microbial biomass carbon content; MBN, microbial biomass nitrogen content; NAG, β-1,4-acetylglucosidase activity; NH4+-N, ammonium nitrogen content; Nmin, nitrogen mineralization rate; NO3--N, nitrate nitrogen content; POX+PER, oxidase activity; SLA, specific leaf area; Soil C:N, soil carbon to nitrogen ratio; SWC, soil water content; TC, total carbon content; TN, nitrogen content. Red represents positive correlation, and blue represents negative correlation. The darker the color, the stronger the correlation. The asterisk indicates a significant correlation (*, p < 0.05; **, p < 0.01; ***, p < 0.001).

图5 方差分解分析探究北京东灵山白桦和蒙古栎林植物根叶功能性状对根际效应变异的解释。A, 白桦。B, 蒙古栎。

Fig. 5 Variations in the rhizosphere effects are partitioned by leaf and root functional traits of Betula platyphylla (A) forest and Quercus mongolica (B) forest in Dongling Mountain, Beijing using variation partitioning analysis.

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