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

Abstract

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

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.

Figures/Tables 11

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References 44

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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

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

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

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

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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

	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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