冬小麦免耕覆盖与有机栽培对土壤微生物群落组成的影响

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王小玲^{1, 2,},
马琨^1,,,
汪志琴^{1, 2},
李越²,
魏常慧²
1.宁夏大学农学院银川 750021
2.宁夏大学西北土地退化与生态恢复国家重点实验室培育基地银川 750021
基金项目: 国家自然科学基金项目31660132
国家自然科学基金项目31160104
宁夏高等学校一流学科建设项目NXYLXK2017B06

详细信息

作者简介:王小玲, 研究方向为农业生态学。E-mail:928176524@qq.com

通讯作者:马琨, 研究方向为农田生态学和土壤微生物生态学。E-mail:makun0411@163.com

中图分类号:S314

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收稿日期:2018-06-24
录用日期:2018-09-30
刊出日期:2019-02-01

Effects of no-tillage, mulching and organic fertilization on soil microbial com-position in winter wheat field

WANG Xiaoling^{1, 2,},
MA Kun^1,,,
WANG Zhiqin^{1, 2},
LI Yue²,
WEI Changhui²
1. College of Agronomy, Ningxia University, Yinchuan 750021, China
2. National Key Laboratory Breeding Base of Northwest Land Degradation and Ecological Restoration, Ningxia University, Yinchuan 750021, China
Funds: the National Natural Science Foundation of China31660132
the National Natural Science Foundation of China31160104
the First-class Discipline Construction Project of Colleges and Universities in NingxiaNXYLXK2017B06

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Corresponding author:MA Kun, E-mail:makun0411@163.com

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摘要
摘要:为揭示农业管理活动对土壤微生物群落结构及AM真菌多样性的影响，以农田土壤生态系统为研究对象，选取免耕覆盖+施有机肥（NF）、免耕覆盖+不施有机肥（NC）、传统耕作不覆盖+施有机肥（TF）和传统耕作不覆盖+不施有机肥（TC）4种处理，采用Illumina Miseq高通量测序及磷脂脂肪酸（phospholipid fatty acids，PLFAs）分析方法，研究持续冬小麦免耕覆盖有机栽培3年后，土壤微生物群落结构组成、AM真菌及其多样性与土壤环境因子间的关系。结果表明，持续免耕覆盖有机栽培能增加以PLFA表征的土壤微生物群落的生物量，传统耕作显著提高了土壤革兰氏阳性菌（G⁺）、阴性菌（G^-）的生物量（P < 0.05）。且随免耕栽培管理年限增加，土壤AM真菌生物量呈显著上升趋势；以16：1ω5c中性脂（NLFA）与16：1ω5c磷脂（PLFA）表征的AM真菌生物量比值显著升高（P < 0.05）。免耕覆盖措施下，有机肥的施用提高了土壤AM真菌丰富度指数（Chao1指数和ACE指数），但降低了土壤AM真菌的多样性（香农指数、辛普森指数）。主成分分析结果显示，AM真菌孢子（16：1ω5c中性脂）的生物量与土壤有机质、土壤易提取球囊霉素含量呈正相关关系，AM真菌丰富度指数与土壤有机质含量呈正相关，AM真菌多样性指数与土壤全氮含量、脲酶活性呈正相关。受农业管理措施导致的土壤理化性状及土壤生物学差异等综合因素影响，土壤微生物生物量及AM真菌多样性和丰富度改变。免耕覆盖措施提高了土壤AM真菌多样性指数，有机肥施用显著影响了AM真菌NLFA/PLFA生物量的比例，改变了AM真菌孢子和菌丝间生物量碳的分配关系。
关键词:免耕覆盖/
有机栽培/
冬小麦/
微生物群落/
AM真菌/
环境因子
Abstract:Soil micro-organisms constitute a significant part of soil fertility and play a critical role in maintaining soil ecological functions. Micro-organisms also are key indicators for soil quality and productivity. No-tillage and mulching cultivation indirectly affect the composition of soil microbial community by changing plant physiological characteristics and root exudates. The two agronomic practices can also improve soil environment by increasing the total amount of micro-organisms in the soil. By analyzing the composition of soil microbial community under no-tillage, mulching and organic cultivation of winter wheat in the southern mountain areas of Ningxia, the effects of agricultural management activities on soil microbial community structure and arbuscular mycorrhizal (AM) fungal diversity were determined in this paper. The study aimed at providing theoretical basis for the promotion of long-term no-tillage, mulching cultivation, sustainable use of farmlands and maintenance of soil microbial diversity. Short-term test on conservation tillage (no-tillage) was done for three consecutive years in Longde County, Guyuan. Four soil treatments were selected including no-tillage and straw mulching without organic fertilizer application (NC), no-tillage and straw mulching with organic fertilizer application (NF), traditional tillage and no-mulching without organic fertilizer application (TC) and traditional tillage and no-mulching with organic fertilizer application (TF). By using Illumina Miseq high-throughput sequencing platform and phospholipid fatty acids (PLFAs) analysis methods, soil microbial community composition, AM fungi community composition and diversity and soil environmental factors were analyzed after three years cultivation of winter wheat. The results showed that continuous no-tillage, mulching and organic fertilizer application increased soil microbial community biomass characterized by PLFA. Traditional tillages significantly improved microbial community biomass (P < 0.05) of gram-positive (G⁺) and negative (G^-) bacteria in the soil. With increasing years of management of no-tillage and straw mulching, soil AM fungi biomass significantly increased. Also biomass ratios of 16:1ω5c neutral lipid (NLFA) to 16:1ω5c phospholipid lipid (PLFA) significantly increased (P < 0.05). Under no-tillage and straw mulching, the application of organic fertilizer increased soil AM fungi richness index (Chao1 index and ACE index), but reduced soil AM fungi diversity (Shannon index and Simpson index). The results of principal component analysis showed that biomass of AM fungal spores (16:1ω5 c neutral fat) was positively correlated with soil contents of organic matter, and easily extracted glomalin. While AM fungi richness index was positively correlated with soil organic matter content, AM fungi diversity index was positively correlated with soil total nitrogen content and urease activity. AM fungi richness and diversity were affected by soil physical and chemical properties and soil biological differences. Also soil microbial biomass changed along with AM fungal diversity and richness. It was concluded that no-tillage and straw mulching cultivation increased soil AM fungi diversity. The proportion of NLFA to PLFA biomass of AM fungi was also significantly affected by the application of organic fertilizer, and changed distribution of biomass carbon between AM fungi spores and mycelium.
Key words:No-tillage and straw mulching/
Organic cultivation/
Winter wheat/
Microbial community/
Arbuscular mycorrhizal fungi (AMF)/
Environmental factors

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图1AM真菌生物量(a)、多样性指数(b)与土壤理化、生物学性状间的多元分析
TN、TP、AP、AN、AK和OM分别指土壤全氮、全磷、有效磷、碱解氮、速效钾和有机质; URE、ALP、CAT、EE-GRSP、T-GRSP分别指脲酶、碱性磷酸酶、过氧化氢酶、易提取球囊霉素、总球囊霉素; AMN、AMP、AMT、NLFA/PLFA分别指AM真菌中性脂、AM真菌磷脂脂肪酸、AM真菌总生物量、中性脂与磷脂脂肪酸的比值; Chao1、ACE、Shannon、Simpson分别指Chao1指数、ACE指数、香浓指数和辛普森指数。图中各形状代表不同采样点。NC表示免耕不施肥的3个重复(1~3), NF表示免耕施肥的3个重复(1~3), TC表示传统耕作不施肥的3个重复(1~3), TF表示传统耕作施肥的3个重复(1~3)。
Figure1.Multivariate analysis of AM fungal biomass (a), AM fungal diversity (b) with soil physical, chemical and biological characteristics
TN, TP, AP, AN, AK and OM refer to total nitrogen, total phosphorus, available phosphorus, available nitrogen, available potassium and organic matter; URE, ALP, CAT, EE-GRSP, T-GRSP refer to urease, alkaline phosphatase, catalase, easily extracted glomalin and total extracted glomalin; AMN, AMP, AMT, NLFA/PLFA refer to AM fungi neutral lipid acid, AM fungi phospholipid acid, AM fungi total biomass, neutral lipid acid and phospholipid acid ratio; Chao1, ACE, Shannon and Simpson refer to Chao 1 index, ACE index, Shannon index, and Simpson index, respectively. Each shape in the figure represents a sample. NC represents three replicates of treatment of no-tillage and straw mulching without organic fertilizer application (1-3), NF represents three replicates of treatment of no-tillage and straw mulching with organic fertilizer application (1-3), and TC represents three replicates of treatment of traditional tillage and no-mulching without organic fertilizer application (1-3), TF shows three replicates of treatment of traditional tillage and no-mulching with organic fertilizer (1-3).

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表1连续免耕覆盖与有机栽培对土壤主要微生物类群的影响
Table1.Effects of continuous no-tillage, straw mulching and organic fertilizer application of winter wheat on soil microorganisms communities

年份 Year	处理 Treatment	细菌 Bacteria (μg?g^-1)	真菌 Fungi (μg?g^-1)	放线菌 Actinomycetes (μg?g^-1)	真菌/细菌 Fungi/bacteria	革兰氏阳性菌 Gram positive bacteria (G⁺)(μg?g^-1)	革兰氏阴性菌 Gram negative bacteria (G^-)(μg?g^-1)	革兰氏阳性菌/革兰氏阴性菌 G⁺/G^-
2015	NC	26.98±2.79a	8.56±1.16b	6.96±1.08a	0.32±0.02ab	8.76±1.07a	18.21±1.91a	0.48±0.04a
	NF	30.53±3.77a	8.77±0.78ab	7.87±0.98a	0.29±0.02b	10.19±2.03a	20.34±1.95a	0.50±0.06a
	TC	27.88±3.60a	8.24±0.95b	7.76±1.39a	0.30±0.01b	9.45±1.31a	18.43±2.34a	0.51±0.02a
	TF	28.94±5.57a	10.25±0.86a	8.50±0.84a	0.36±0.05a	10.14±1.18a	18.88±4.74a	0.54±0.14a
2017	NC	31.67±2.74ab	6.29±0.48b	7.59±0.38ab	0.20±0.20a	14.43±1.37ab	17.24±1.68b	0.84±0.07ab
	NF	29.59±5.90b	6.63±1.68ab	7.35±1.52b	0.22±0.01a	12.99±2.55b	16.60±3.38b	0.78±0.03bc
	TC	32.22±3.80ab	6.47±0.38ab	7.50±0.26ab	0.20±0.02a	15.18±2.79a	17.04±1.55b	0.89±0.13a
	TF	36.21±5.60a	7.75±1.18a	8.48±0.91a	0.22±0.01a	15.62±3.39a	20.59±2.61a	0.76±0.07c
不同小写字母代表不同处理间差异在0.05水平上显著。NC:免耕覆盖不施有机肥; NF:免耕覆盖施有机肥; TC:传统耕作不覆盖不施有机肥; TF:传统耕作不覆盖施有机肥。Different lowercase letters indicate significant differences among treatments at 0.05 level. NC: no-tillage and straw mulching without organic fertilizer application; NF: no-tillage and straw mulching with organic fertilizer application; TC: traditional tillage and no-mulching without organic fertilizer application; TF: traditional tillage and no-mulching with organic fertilizer application.

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表2连续免耕覆盖有机管理对土壤AM真菌群落结构组成的影响
Table2.Effects of no-tillage, straw mulching and organic fertilizer application of winter wheat on soil AM fungal community structure

年份 Year	处理 Treatment	AM真菌磷脂脂肪酸 AMF PLFAs (μg·g^-1)	AM真菌中性脂 AMF NLFA (μg·g^-1)	AM真菌总生物量 Total AMF (μg·g^-1)	中性脂/磷脂脂肪酸 NLFA / PLFA	微生物总磷脂脂肪酸 Total PLFAs (μg·g^-1)	AM真菌/微生物总磷脂脂肪酸 AMF / total PLFAs
2015	NC	2.20±0.24a	0.87±0.01a	3.07±0.66a	0.40±0.00a	45.23±5.22a	0.07±0.10a
	NF	2.47±0.35a	0.76±0.38a	3.23±0.72a	0.30±0.11a	50.74±6.02a	0.06±0.01a
	TC	2.38±0.30a	1.24±1.03a	3.62±1.25a	0.52±0.39a	47.83±6.94a	0.08±0.07a
	TF	2.59±0.21a	0.60±0.33a	3.19±0.44a	0.23±0.13a	51.25±6.85a	0.07±0.01a
2017	NC	2.17±0.19ab	16.84±4.53a	19.01±4.65a	7.72±1.74a	64.56±6.21a	0.29±0.50a
	NF	2.13±0.15b	7.62±4.47b	9.75±4.35b	3.64±2.28b	53.32±5.61b	0.19±1.28a
	TC	2.15±0.26ab	6.66±4.94b	8.81±5.12b	3.03±1.97b	54.99±5.95b	0.16±2.01a
	TF	2.45±0.34a	9.54±7.08ab	11.99±6.96b	4.01±3.41b	65.37±9.91a	0.18±2.70a
不同小写字母代表不同处理间差异在0.05水平上显著。NC:免耕覆盖不施有机肥; NF:免耕覆盖施有机肥; TC:传统耕作不覆盖不施有机肥; TF:传统耕作不覆盖施有机肥。Different lowercase letters indicate significant differences among treatments at 0.05 level. NLFA: neutral lipid acid. NC: no-tillage and straw mulching without organic fertilizer application; NF: no-tillage and straw mulching with organic fertilizer application; TC: traditional tillage and no-mulching without organic fertilizer application; TF: traditional tillage and no-mulching with organic fertilizer application.

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表3冬小麦免耕栽培对土壤AM真菌丰富度和多样性指数的影响
Table3.Effects of no-tillage, straw mulching and organic application of winter wheat on soil AM fungal richness and diversity

处理 Treatment	辛普森指数 Simpson index	香农指数 Shannon index	Chao1指数 Chao1 index	ACE指数 ACE index
NC	0.95±0.02a	5.95±0.45a	668.99±99.77a	696.34±127.06a
NF	0.93±0.03a	5.45±0.38a	682.03±166.58a	714.67±195.47a
TC	0.92±0.03a	5.45±0.26a	702.53±67.55a	754.21±81.15a
TF	0.89±0.05a	5.25±0.67a	742.02±178.85a	783.18±178.96a
不同小写字母代表不同处理间差异在0.05水平上显著。NC:免耕覆盖不施有机肥; NF:免耕覆盖施有机肥; TC:传统耕作不覆盖不施有机肥; TF:传统耕作不覆盖施有机肥。Different lowercase letters indicate significant differences among treatments at 0.05 level. NC: no-tillage and straw mulching without organic fertilizer application; NF: no-tillage and straw mulching with organic fertilizer application; TC: traditional tillage and no-mulching without organic fertilizer application; TF: traditional tillage and no-mulching with organic fertilizer application.

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表4Mantel检验土壤AM真菌群落结构及AM真菌多样性与土壤理化、生物学性状间的关系
Table4.Correlation between community structure and diversity of AM fungi and soil physical and chemical biological characteristics

变量 Variable	中性脂/磷脂脂肪酸 NLFA /PLFA	AM真菌磷脂脂肪酸 AMF PLFA	AM真菌中性脂 AMF NLFA	AM真菌生物量 Total AMF	辛普森指数 Simpson index	香农指数 Shannon index	ACE指数 ACE index	Chao1指数 Chao 1 index
土壤有机质Soil organic matter	-0.01	0.51	0.07	0.09	-0.44	-0.12	0.57	0.62
全氮Total nitrogen	0.61	-0.68	0.57	0.55	0.85	0.89^*	-0.77	-0.69
全磷Total phosphorus	-0.32	0.48	-0.24	-0.23	-0.60	-0.38	0.68	0.78
碱解氮Available nitrogen	0.50	-0.77	0.45	0.42	0.85	0.83	-0.79	-0.68
速效钾Available potassium	-0.05	-0.94^*	-0.12	-0.15	0.68	0.43	-0.71	-0.54
pH	0.52	0.22	0.57	0.58	0.08	0.44	0.08	0.10
速效磷Available phosphorus	0.08	-0.33	0.02	0.01	0.35	0.10	-0.47	-0.57
脲酶Urease	-0.53	0.87	-0.46	-0.44	-0.96^**	-0.87	0.93^*	0.86
过氧化氢酶Catalase	0.17	-0.83	0.07	0.05	0.77	0.45	-0.86	-0.85
碱性磷酸酶Alkaline phosphatase	-0.70	0.37	-0.69	-0.68	-0.65	-0.84	0.52	0.47
易提取球囊霉素Easily extracted glomalin	0.86	0.42	0.91^*	0.92^*	0.16	0.54	-0.02	-0.17
总球囊霉素Total extracted glomalin	0.79	0.52	0.84	0.85	0.05	0.41	0.05	-0.13
^和^分别代表 0.05和0.01水平显著相关。^ and ^** respectively represent significant correlations at 0.05 and 0.01 levels. NLFA: neutral lipid acid.

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