黄土高原生物结皮形成过程中土壤胞外酶活性及其化学计量变化特征

doi:10.13866/j.azr.2022.02.13

干旱区研究 ›› 2022, Vol. 39 ›› Issue (2): 456-468.doi: 10.13866/j.azr.2022.02.13

黄土高原生物结皮形成过程中土壤胞外酶活性及其化学计量变化特征

姚宏佳¹(),王宝荣^2,³,安韶山^1,²,杨娥女¹,黄懿梅⁴

1.西北农林科技大学水土保持研究所,黄土高原土壤侵蚀与旱地农业国家重点实验室,陕西杨凌 712100
2.中国科学院水利部水土保持研究所,黄土高原土壤侵蚀与旱地农业国家重点实验室,陕西杨凌 712100
3.中国科学院大学,北京 100049
4.西北农林科技大学资源与环境学院,农业部西北植物营养与农业环境重点实验室,陕西杨凌 712100

收稿日期:2021-08-24 修回日期:2021-11-24 出版日期:2022-03-15 发布日期:2022-03-30
作者简介:姚宏佳（1997-）, 女, 硕士研究生, 主要从事土壤微生物生态研究. E-mail: yaohj97@163.com
基金资助:
国家自然科学基金面上项目(41671280);国家自然科学基金面上项目(41877074);国家自然科学基金面上项目(42077072)

Variation in soil extracellular enzyme activities stoichiometry during biological soil crust formation in the Loess Plateau

YAO Hongjia¹(),WANG Baorong^2,³,AN Shaoshan^1,²,YANG E’nv¹,HUANG Yimei⁴

1. State Key Laboratory of Soil Erosion and Dryland Farming on the Loess Plateau, Institute of Soil and Water Conservation, Northwest A & F University, Yangling 712100, Shaanxi, China
2. State Key Laboratory of Soil Erosion and Dryland Farming on the Loess Plateau, Institute of Soil and Water Conservation, Chinese Academy of Science and Ministry of Water Resources, Yangling 712100, Shaanxi, China
3. University of Chinese Academy of Sciences, Beijing 100049, China
4. Key Laboratory of Plant Nutrition and the Agro-Environment in Northwest China, Ministry of Agriculture, College of Natural Resources and Environment, Northwest A & F University, Yangling 712100, Shaanxi, China

Received:2021-08-24 Revised:2021-11-24 Online:2022-03-15 Published:2022-03-30

摘要/Abstract

摘要：

生物土壤结皮在增强土壤抵抗水蚀风蚀能力、改善土壤养分等方面发挥着重要作用,土壤胞外酶活性可作为土壤生化反应强度的微生物指标,对理解荒漠生态系统微生物参与的养分循环过程具有重要意义。选取黄土高原神木县六道沟小流域5个生物结皮发育阶段（裸沙地、全藻结皮、藻-藓混生结皮、藓-藻混生结皮、全藓结皮）的土壤作为研究对象,研究生物结皮发育过程中土壤胞外酶活性（β-1,4-葡萄糖苷酶、β-1,4-N-乙酰基氨基葡萄糖苷酶、亮氨酸氨基多肽酶和碱性磷酸酶）及其化学计量的变化特征。结果表明：（1）土壤4种胞外酶活性均随生物结皮发育序列变化显著增加,全藓结皮土壤显著大于全藻结皮土壤（P<0.05）。（2）生物土壤结皮层胞外酶活性显著高于下层土壤,且胞外酶活性随着土层深度的加深不断降低。（3）相关性分析表明,土壤C、N、P、土壤C：P、土壤N：P均与土壤胞外酶活性呈极显著正相关关系（P<0.05）。（4）有结皮覆盖土壤的养分含量及土壤胞外酶活性显著高于裸沙地,全藓结皮土壤的养分含量及土壤胞外酶活性显著高于全藻结皮土壤。（5）标准化主轴估计表明,土壤胞外酶活性随着生物结皮的发育均有明显增强并呈现稳态特征,N-获取酶和P-获取酶相对于C-获取酶之间的斜率呈现等容关系,这一结果表明土壤微生物通过稳态调控胞外酶化学计量特征,在干旱半干旱生态系统养分循环过程中具有重要作用。

关键词: 生物土壤结皮, 发育序列, 胞外酶, 化学计量, 黄土高原

Abstract:

Biological soil crusts (BSCs) play an important role in enhancing soil resistance to water and wind erosion, and improving soil nutrients. Soil extracellular enzyme activity can be used as a microbial indicator of the intensity of soil biochemical reactions, which is important for understanding the nutrient cycling processes involving microorganisms in desert ecosystems. However, there is a lack of understanding of the variation in soil extracellular enzymes and their stoichiometry during the formation of BSCs on the Loess Plateau. Soils from five biological crust formation stages (CK, A, AM, MA, and M) in Liudaogou watershed, Shenmu County, on the Loess Plateau were selected for the study. Soil extracellular enzyme activities [β-1,4-glucosidase (BG), β-1,4-N-acetylaminoglucosidase (NAG), leucine aminopeptidase (LAP), and alkaline phosphatase (AP)] and their stoichiometric changes during the formation of biocrusts were characterized. The activities of soil BG, NAG, LAP, and AP increased significantly with the sequence of biological crust development, and activity was significantly greater in Mo soil than that in Al soil (P<0.05). Extracellular enzyme activities were significantly higher in the BSCs layer than those in the lower soil layer, and decreased continuously with the depth of the soil. Soil C, N, P, C:P, and N:P all showed highly significant positive correlations with soil extracellular enzyme activities (P<0.05). The nutrient content and soil extracellular enzyme activity of soil with crust cover were significantly higher than those of bare land, and those of Mo soil were significantly higher than those of Al soil. Standardized major axis estimates indicated that soil extracellular enzyme activities were significantly enhanced and showed homeostasis with formation of biological crusts, and the slopes between N- and P-acquiring enzymes relative to C-acquiring enzymes showed isotropic relationships. Soil microorganisms play an important role in nutrient cycling in arid and semi-arid ecosystems through homeostatic regulation of extracellular enzyme stoichiometry.

Key words: biological soil crusts, developmental sequence, extracellular enzymes, stoichiometry, Loess Plateau

姚宏佳,王宝荣,安韶山,杨娥女,黄懿梅. 黄土高原生物结皮形成过程中土壤胞外酶活性及其化学计量变化特征[J]. 干旱区研究, 2022, 39(2): 456-468.

YAO Hongjia,WANG Baorong,AN Shaoshan,YANG E’nv,HUANG Yimei. Variation in soil extracellular enzyme activities stoichiometry during biological soil crust formation in the Loess Plateau[J]. Arid Zone Research, 2022, 39(2): 456-468.

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发育阶段	纬度	经度	海拔/m
裸沙地CK	38°47′3.98″	110°21′44.57″	1207
全藻结皮A	38°47′7.45″	110°21′32.99″	1224
藻-藓混生结皮AM	38°46′57.49″	110°21′29.58″	1215
藓-藻混生结皮MA	38°46′51.58″	110°21′25.82″	1210
全藓结皮M	38°45′57.64″	110°21′6.86″	1242

养分	土层/cm	发育阶段
养分	土层/cm	CK	A	AM	M	M
SOC/(g·kg ^-1)	BSCs		15.2±3.08Ba	17.8±3.20ABa	20.0±2.20Aa	19.8±3.97Aa
	0~2	2.44±0.29Ba	6.73±1.97Ab	5.45±1.24Ab	5.80±1.97Ab	6.14±2.76Ab
	2~10	1.53±0.37Bb	2.44±0.65Ac	2.27±0.46ABc	2.09±0.69ABc	2.41±0.69Ac
	10~20	1.29±0.28Ab	1.28±0.43Ac	1.72±0.44Ac	1.61±0.37Ac	1.47±0.47Ac
TN/(g·kg ^-1)	BSCs		1.10±0.27Ba	1.15±0.23Ba	1.38±0.16Aa	1.30±0.26ABa
	0~2	0.10±0.01Ba	0.46±0.14Ab	0.42±0.11Ab	0.49±0.14Ab	0.46±0.19Ab
	2~10	0.09±0.02Bb	0.18±0.05Ac	0.18±0.05Ac	0.16±0.05Ac	0.17±0.05Ac
	10~20	0.08±0.01Bc	0.10±0.03ABc	0.13±0.04ABc	0.13±0.02Ac	0.11±0.04Ac
TP/(g·kg ^-1)	BSCs		0.40±0.03Ba	0.40±0.04ABa	0.45±0.06Aa	0.39±0.03Ba
	0~2	0.19±0.02Ba	0.32±0.03Ab	0.32±0.04Ab	0.34±0.05Ab	0.31±0.03Ab
	2~10	0.16±0.03Bb	0.24±0.09ABc	0.27±0.12Ab	0.23±0.03ABc	0.23±0.02ABc
	10~20	0.16±0.01Bb	0.19±0.06ABc	0.20±0.02Ac	0.20±0.03Ac	0.19±0.02ABd

胞外酶	土层 /cm	发育阶段
胞外酶	土层 /cm	CK	A	Mo	MA	M
BG/(nmol·h^-1·g^-1)	BSCs		80.1±10.5ABa	71.6±9.81Ba	117±6.89Aa	111±19.3Aa
	0~2	19.5±1.65Ba	41.5±6.83ABb	26.9±5.11ABb	42.6±5.68Ab	36.5±12.5ABb
	2~10	13.7±1.95Ab	15.9±3.21Ac	12.2±2.51Ab	12.3±1.70Ac	13.6±3.14Ab
	10~20	8.71±1.13BCc	6.51±1.47BCc	10.1±1.98ABb	13.5±2.02Ac	4.67±0.85Cc
NAG/(nmol·h^-1·g^-1)	BSCs		19.3±3.35Ba	23.9±2.15Ba	65.1±4.80Aa	61.0±8.45Aa
	0~2	3.75±0.37Bb	7.13±1.41ABb	5.14±0.70Bb	12.3±1.60Ab	12.5±3.90Ab
	2~10	11.6±1.70Aa	7.04±1.43Ab	6.79±1.55Ab	9.68±1.74Ab	8.68±1.73Ab
	10~20	10.8±2.97Aa	8.16±1.38Ab	8.80±1.55Ab	9.85±1.26Ab	9.99±2.12Ab
LAP/(nmol·h^-1·g^-1)	BSCs		167±13.5Aa	150±25.7Aa	145±12.3Aa	206±36.8Aa
	0~2	11.3±0.76Ca	67.7±5.23Ab	60.8±8.08Bb	86.4±6.77Ab	86.2±10.31Ab
	2~10	8.85±0.63Bb	40.0±5.17Ac	42.4±7.69Ab	36.1±6.17Ac	49.0±7.81Abc
	10~20	10.3±0.72Cab	20.0±2.38BCc	19.0±2.41ABCb	13.7±3.59ABc	23.9±4.65Ac
AP/(nmol·h^-1·g^-1)	BSCs		80.0±16.2Ba	72.6±5.75Ba	143±5.56Aa	132±5.04Aa
	0~2	11.9±1.24Ba	18.5±2.13Bb	11.7±1.58Bb	35.1±2.66Ab	31.9±4.67Ab
	2~10	10.3±1.91Bab	9.88±1.09Bb	14.9±4.06Bb	29.3±4.37ABb	20.2±5.92Ab
	10~20	6.18±0.98Bb	10.9±1.07Bb	9.90±1.79Bb	28.9±4.28Ab	22.8±4.83Ab

发育阶段	变量		R²	P	斜率	P(test)
发育阶段	X	Y	R²	P	斜率	P(test)
裸沙地CK	BG	NAG+LAP	0.055	0.211	0.673	0.035
	BG	AP	0.168	<0.05	1.128	0.489
	NAG+LAP	AP	0.000	0.960	/	/
全藻结皮A	BG	NAG+LAP	0.682	<0.001	0.675	0.000
	BG	AP	0.372	<0.001	0.784	0.064
	NAG+LAP	AP	0.681	<0.001	1.162	0.107
藻-藓混生结皮AM	BG	NAG+LAP	0.272	<0.01	0.715	0.018
	BG	AP	0.501	<0.001	1.066	0.578
	NAG+LAP	AP	0.287	<0.001	1.491	0.005
藓-藻混生结皮MA	BG	NAG+LAP	0.667	<0.001	0.849	0.087
	BG	AP	0.647	<0.001	0.771	0.010
	NAG+LAP	AP	0.598	<0.001	0.908	0.352
全藓结皮M	BG	NAG+LAP	0.554	<0.001	0.501	0.000
	BG	AP	0.244	<0.01	0.597	0.000
	NAG+LAP	AP	0.445	<0.001	1.192	0.151
总计	SOC	BG	0.618	<0.001	1.116	0.016
	TN	NAG+LAP	0.797	<0.001	0.911	0.005
	TP	AP	0.418	<0.001	2.978	0.000

组分	发育阶段(S)		土层(L)/cm		S×L
组分	F	P	F	P	F	P
C	6.581	0.000^***	700.6	0.000^***	4.982	0.000^***
N	10.938	0.000^***	592.809	0.000^***	4.393	0.000^***
P	19.868	0.000^***	149.354	0.000^***	2.258	0.014^*
C：N	46.178	0.000^***	10.84	0.000^***	5.265	0.000^***
C：P	2.434	0.049^*	433.542	0.000^***	3.751	0.000^***
N：P	9.646	0.000^***	443.635	0.000^***	4.044	0.000^***
BG	3.432	0.010^*	127.296	0.000^***	2.46	0.007^**
NAG	18.582	0.000^***	125.371	0.000^***	11.751	0.000^***
LAP	7.249	0.000^***	111.851	0.000^***	1.729	0.071
NAG+LAP	7.962	0.000^***	139.847	0.000^***	2.337	0.011^*
AP	30.830	0.000^***	282.224	0.000^***	5.772	0.000^***
BG：（NAG+LAP）	13.282	0.000^***	10.033	0.000^***	3.563	0.000^***
BG:AP	7.637	0.000^***	7.062	0.000^***	1.168	0.313
（NAG+LAP）：AP	4.496	0.002^**	3.456	0.018^*	1.27	0.246

黄土高原生物结皮形成过程中土壤胞外酶活性及其化学计量变化特征

Variation in soil extracellular enzyme activities stoichiometry during biological soil crust formation in the Loess Plateau

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土壤胞外酶	缩写	对应底物
β-1,4-葡萄糖苷酶	BG	4-MUB-β-D-glucoside
β-1,4-乙酰基氨基葡萄糖苷酶	NAG	4-MUB-N-acetyl-β-D-glucosaminide
亮氨酸氨基多肽酶	LAP	L-leucine-7-amido-4-methylcoumarin hydrochloride
碱性磷酸酶	AP	4-MUB-phosphate

	C	N	P	C:N	C:P	N:P	BG	NAG	LAP	NAG+LAP	AP	BG:(NAG+LAP)	BG:AP	(NAG+LAP):AP
C	1
N	0.981^**	1
P	0.806^**	0.834^**	1
C：N	0.074	-0.053	-0.167^*	1
C：P	0.981^**	0.946^**	0.725^**	0.159^*	1
N：P	0.970^**	0.984^**	0.776^**	-0.049	0.965^**	1
BG	0.859^**	0.878^**	0.719^**	-0.019	0.831^**	0.868	1
NAG	0.792^**	0.784^**	0.575^**	0.005	0.756^**	0.760^**	0.853^**	1
LAP	0.808^**	0.808^**	0.718^**	-0.058	0.789^**	0.805^**	0.788^**	0.667^**	1
NAG+LAP	0.855^**	0.853^**	0.729^**	-0.047	0.830^**	0.845^**	0.851^**	0.785^**	0.985^**	1
AP	0.813^**	0.808^**	0.665^**	-0.001	0.778^**	0.787^**	0.745^**	0.762^**	0.707^**	0.763^**	1
BG：(NAG+LAP)	0.078	0.083	0.001	0.337^**	0.112	0.098	0.243^**	0.067	-0.182^*	-0.136	0.007	1
BG：AP	-0.032	-0.018	0.089	0.09	-0.021	-0.011	0.176^*	-0.032	-0.029	-0.032	-0.281^**	0.442^**	1
(NAG+LAP)：AP	-0.115	-0.111	-0.044	-0.068	-0.119	-0.113	-0.109	-0.126	0.049	0.012	-0.221^**	-0.208^**	0.321^**	1

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