长期保护性耕作对丛枝菌根真菌多样性的影响

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张贵云^1,,
张丽萍^1,,,
魏明峰¹,
刘珍¹,
范巧兰¹,
吕贝贝¹,
姚众¹,
柴跃进²
1.山西省农业科学院棉花研究所运城 044000
2.山西省临汾市农机局临汾 041000
基金项目: 山西省自然科学基金项目2014011029-3
山西省农业科学院博士基金项目YBSJJ1405
山西省重点研发计划项目201703D321009-3

详细信息

作者简介:张贵云, 主要从事植物营养学和菌根学研究。E-mail:guiyunzhang@126.com

通讯作者:张丽萍, 主要从事农作物病虫害综合治理和植物营养学研究。E-mail:lipingzh2006@126.com

中图分类号:S34

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收稿日期:2017-12-01
录用日期:2018-02-14
刊出日期:2018-07-01

Effect of long-term conservation tillage on arbuscular mycorrhizal fungi diversity

ZHANG Guiyun^1,,
ZHANG Liping^1,,,
WEI Mingfeng¹,
LIU Zhen¹,
FAN Qiaolan¹,
LYU Beibei¹,
YAO Zhong¹,
CHAI Yuejin²
1. Cotton Research Institute, Shanxi Academy of Agricultural Sciences, Yuncheng 044000, China
2. Agricultural Machinery Bureau in Linfen City of Shanxi Province, Linfen 041000, China
Funds: the Natural Science Foundation of Shanxi Provicne, China2014011029-3
the Doctor Fund Project of Shanxi Academy of Agricultural Sciences, ChinaYBSJJ1405
the Key Research and Development Project of Shanxi Provicne, China201703D321009-3

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Corresponding author:ZHANG Liping, E-mail: lipingzh2006@126.com

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摘要
摘要:为了明确我国北方干旱地区长期保护性耕作以及深松对丛枝菌根真菌（AMF）多样性的影响，笔者于2014年在山西省临汾市连续22年实施保护性耕作的长期定位试验基地，针对免耕覆盖（NTS）、深松免耕覆盖（SNTS）及传统耕作（TT）3种处理方式，进行了不同耕作条件下土壤AMF物种丰度、孢子密度、Shannon多样性指数以及AMF侵染率等因素的比较研究。结果显示，长期保护性耕作（NTS和SNTS）共分离鉴定出AMF 7属9种，其中根孢囊霉属（Rhizophagus）和斗管囊霉属（Funneliformis）各2种，球囊霉属（Glomus）、近明球囊霉属（Claroideoglomus）、无梗囊霉属（Acaulospora）、硬囊霉属（Sclerocystis）和隔球囊霉属（Septoglomus）各1种；而传统耕作（TT）共分离鉴定出AMF 6属8种，没有检测到无梗囊霉属。NTS、SNTS和TT处理在不同土层的AMF优势种基本一致，0~40 cm土层为摩西斗管囊霉（Fu.mosseae）和变形球囊霉（G.versiforme），40~80 cm土层为摩西斗管囊霉、变形球囊霉和聚丛根孢囊霉（Rh.aggregatum），80~120 cm土层为聚丛根孢囊霉，120 cm土层以下只有NTS和SNTS处理中存在聚丛根孢囊霉，说明保护性耕作措施促进了AMF向土壤深层发展。NTS和SNTS处理在同一土层的AMF物种丰度、孢子密度和Shannon多样性指数均高于TT处理，SNTS处理高于NTS处理。同一耕作措施不同土层的AMF物种丰度、孢子密度和Shannon多样性指数均随土层加深而逐渐降低；NTS和SNTS处理在小麦各生育期的丛枝侵染率和孢子密度均高于TT处理；各处理在小麦拔节期的AMF侵染率最高，分别为14.9%、16.1%和10.6%，而在收获期的土壤孢子密度最高，分别为111.7个·（100g）^-1、125.0个·（100g）^-1和90.3个·（100g）^-1。研究认为，长期免耕覆盖、尤其深松免耕覆盖，提高了AMF多样性。该研究结果可为中国北方旱作农田生态系统中AMF自然潜力的充分发挥，以及保护性耕作技术的合理应用提供科学依据。
关键词:丛枝菌根真菌(AMF)/
免耕深松/
物种丰度/
多样性/
侵染率/
孢子密度
Abstract:To verify the effect of long-term conservation tillage and subsoiling on arbuscular mycorrhizal fungi (AMF) diversity in dry areas in northern China, a comparative analysis of species richness, spore density, Shannon diversity index and colonization rate of AMF among no-till with straw mulching (NTS), no-till with straw mulching after subsoiling (SNTS) and traditional tillage treatment (TT) were conducted in 2014 at a long-term (22 years) positioning test base for conservation tillage in Linfen, Shanxi Province. In the study, 9 AMF species belonging to 7 genera were isolated and identified in soil samples of NTS and SNTS treatments, including two species each of Rhizophagus and Funneliformis and one species each of Glomus, Claroideoglomus, Acaulospora, Sclerocystis and Septoglomus. Also 8 AMF species belonging to 6 genera were isolated and identified in soil samples of TT treatment. The AMF species in the TT treatment were the same with those in NTS and SNTS treatments, except that there was no Acaulospora.The dominant species of AMF in NTS, SNTS and TT treatments were the same. The dominant species of AMF in the 0-40 cm soil layer were mainly Fu. mosseae and G. versiforme, the 40-80 cm soil layer were Fu. mosseae, G. versiforme and Rh. aggregatum, and the 80-120 cm soil layer was Rh. aggregatum. Below the 120 cm soil layer, the dominant AMF species was Rh. aggregatum, which was found only under NTS and SNTS treatments. Species richness, spore density and Shannon diversity index of AMF in NTS and SNTS treatments in the same soil layers were higher than those in TT treatment, and those in SNTS treatment were higher than those in NTS treatment. Species richness, spore density and Shannon diversity index of AMF decreased with increasing of soil depth under the same treatment. The maximum species richness, spore density and Shannon diversity index of AMF were all in the 0-20 cm shallow soil layer. The colonization rate and spore density of AMF at different growth stages of wheat in NTS and SNTS treatments were all significantly higher than that in TT treatment. Colonization rate of AMF in NTS, SNTS and TT treatments was the highest at jointing stage of wheat and was respectively 14.9%, 16.1% and 10.6%. AMF spore density of soil was the highest at maturity stage of wheat and was respectively 111.7 spores·(100g)^-1, 125.0 spores·(100g)^-1 and 90.3 spores·(100g)^-1. The study showed that long-term no-till with straw mulching improved AM fungi diversity, especially under no-till with straw mulching after subsoiling. The results provided the scientific basis for the full play of AMF role and reasonable application of conservation tillage in dry farmland ecosystems in northern China.
Key words:Arbuscular mycorrhizal fungi (AMF)/
No-till with straw mulching after subsoiling/
AMF species richness/
AMF diversity/
Colonization rate/
Spore density

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表1不同耕作方式土壤的理化性状
Table1.Soil physical-chemical characteristics under different tillage treatments

处理 Treatment	pH	有机质 Organic matter (g·kg^-1)	全氮 Total N (g·kg^-1)	碱解氮 Alkali-hydrolysis N (mg·kg^-1)	速效磷 Available P (mg·kg^-1)	速效钾 Available K (mg·kg^-1)	土壤容重 Soil bulk density (g·cm^-3)
处理 Treatment	pH	有机质 Organic matter (g·kg^-1)	全氮 Total N (g·kg^-1)	碱解氮 Alkali-hydrolysis N (mg·kg^-1)	速效磷 Available P (mg·kg^-1)	速效钾 Available K (mg·kg^-1)	0~10 cm	10~20 cm
TT	8.48±0.02a	14.56±0.63b	0.75±0.01b	57.74±2.40b	49.34±1.48b	214.35±9.82b	1.29±0.31	1.35±0.40
NTS	8.35±0.01b	18.23±0.43b	1.02±0.01a	91.00±7.00a	63.60±3.28a	362.73±17.66a	1.21±0.26	1.47±0.42
SNTS	8.36±0.01b	23.82±1.76a	1.10±0.03a	86.75±2.83a	74.21±1.20a	318.16±15.22a	1.19±0.22	1.45±0.42
??NTS:免耕覆盖; SNTS:深松免耕覆盖; TT:传统耕作。同列数字后不同小写字母表示处理间差异显著(P < 0.05)。TT: traditional tillage; NTS: no-till with straw mulching; SNTS: no-till with straw mulching after subsoiling. Different lowercase letters in the same column indicate significant differences at 0.05 level among different treatments.

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表2小麦成熟期不同耕作措施下不同土层丛枝菌根真菌(AMF)种类分布及相对多度
Table2.Species distribution and relative abundance of arbuscular mycorrhizal fungi (AMF) in different soil depths at wheat maturation period under different tillage treatments

%
土层深度 Soil depth (cm)	无梗囊霉属 Acaulospora			近明球囊霉属 Claroideoglomus			斗管囊霉属 Funneliformis						球囊霉属 Glomus			根孢囊霉属 Rhizophagus						硬囊霉属 Sclerocystis			隔球囊霉属 Septoglomus
	细凹无梗囊霉 Ac. scrobiculata			幼套近明球囊霉 Cl. etunicatum			摩西斗管囊霉 Fu. mosseae			地斗管囊霉 Fu. geosporus			变形球囊霉 G. versiforme			聚丛根孢囊霉 Rh. aggregatum			根内根孢囊霉 Rh. intraradices			弯丝硬囊霉 Sc. sinuosus			缩隔球囊霉 Se. constrictum
	NTS	SNTS	TT	NTS	SNTS	TT	NTS	SNTS	TT	NTS	SNTS	TT	NTS	SNTS	TT	NTS	SNTS	TT	NTS	SNTS	TT	NTS	SNTS	TT	NTS	SNTS	TT
0~20	0.3bB	0.5cA	0.0aC	9.8aA	8.8aB	6.9aC	32.2aA	31.3aA	32.0aA	3.4aB	3.6bB	5.6bA	30.3aA	27.2aB	31.4aA	10.1dAB	12.8eA	7.8cB	10.4aB	9.3aC	12.4aA	0.6bB	1.4cA	0.2aC	2.9bC	5.1aA	3.7aB
20~40	0.3bB	0.5cA	0.0aC	9.8aA	8.8aB	6.9aC	32.2aA	31.3aA	32.0aA	3.4aB	3.6bB	5.6bA	30.3aA	27.2aB	31.4aA	10.1dAB	12.8eA	7.8cB	10.4aB	9.3aC	12.4aA	0.6bB	1.4cA	0.0bC	2.9bC	5.1aA	3.7aB
40~60	0.6aB	1.7aA	0.0aC	7.2bA	3.8bC	5.6bB	24.3bA	24.6bA	25.7bA	4.0aB	6.1aA	6.5aC	24.1bA	23.9bA	19.5bB	25.8cA	27.9dA	32.4bB	8.3bA	8.1bA	7.3bB	1.3aB	3.0aA	0.0bC	4.4aA	4.1bA	3.0bB
60~80	0.0cB	1.2bA	0.0aB	7.2bA	2.7cC	5.6bB	24.3bA	17.7cB	25.7bA	4.0aA	4.4bA	6.5aB	24.1bA	23.2bA	19.5bB	25.8cB	39.8cA	32.4bC	8.3bA	5.8cB	7.3bC	1.3aB	2.2bA	0.0bC	4.4aA	3.0cB	3.0bC
80~100	0.0cA	0.0dA	0.0aA	0.0cA	0.0dA	0.0cA	18.3cB	16.5cB	24.0bA	0.0bA	0.0cA	0.0cA	0.0c B	8.5cA	0.0c B	72.4bA	66.0bB	76.0aA	9.3bA	9.0aA	0.0cB	0.0cA	0.0dA	0.0bA	0.0cA	0.0dA	0.0cA
100~120	0.0cA	0.0dA	0.0aA	0.0cA	0.0dA	0.0cA	18.3cB	16.5cB	25.0bA	0.0bA	0.0cA	0.0cA	0.0c B	8.5cA	0.0c B	72.3bB	66.0bC	75.0aA	9.3bA	9.0aA	0.0cB	0.0cA	0.0dA	0.0bA	0.0cA	0.0dA	0.0cA
120~140	0.0cA	0.0dA	0.0aA	0.0cA	0.0dA	0.0cA	0.0dA	0.0dA	0.0cA	0.0bA	0.0cA	0.0cA	0.0cA	0.0dA	0.0cA	100.0aA	100.0aA	0.0aB	0.0cA	0.0dA	0.0cA	0.0cA	0.0dA	0.0bA	0.0cA	0.0dA	0.0cA
140~160	0.0cA	0.0dA	0.0aA	0.0cA	0.0dA	0.0cA	0.0dA	0.0dA	0.0cA	0.0bA	0.0cA	0.0cA	0.0cA	0.0dA	0.0cA	0.0eB	100.0aA	0.0dB	0.0cA	0.0dA	0.0cA	0.0cA	0.0dA	0.0bA	0.0cA	0.0dA	0.0cA
160~180	0.0cA	0.0dA	0.0aA	0.0cA	0.0dA	0.0cA	0.0dA	0.0dA	0.0cA	0.0bA	0.0cA	0.0cA	0.0cA	0.0dA	0.0cA	0.0eA	0.0fA	0.0dA	0.0cA	0.0dA	0.0cA	0.0cA	0.0dA	0.0bA	0.0cA	0.0dA	0.0cA
180~200	0.0cA	0.0dA	0.0aA	0.0cA	0.0dA	0.0cA	0.0dA	0.0dA	0.0cA	0.0bA	0.0cA	0.0cA	0.0cA	0.0dA	0.0cA	0.0eA	0.0fA	0.0dA	0.0cA	0.0dA	0.0cA	0.0cA	0.0dA	0.0bA	0.0cA	0.0dA	0.0cA
??NTS:免耕覆盖; SNTS:深松免耕覆盖; TT:传统耕作。同列不同小写字母表示不同土层间差异显著(P < 0.05);同行不同大写字母表示同一土层不同处理间差异显著(P < 0.05)。NTS: no-till with straw mulching; SNTS: no-till with straw mulching after subsoiling; TT: traditional tillage. Different small letters in the same column mean significant differences among different soil layers in the same treatment at P < 0.05, and different capital letters in the same row mean significant differences in the same soil layer among different treatments at P < 0.05.

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表3小麦成熟期不同耕作措施下丛枝菌根真菌(AMF)物种丰度、孢子密度和多样性指数
Table3.Species richness, spore density and diversity index of arbuscular mycorrhizal fungi (AMF) in different soil depths at wheat maturation period under different tillage treatments

土样深度 Soil depth (cm)	物种丰度 Species richness			孢子密度 Spore density [spores·(100g)^-1(soil)]			Shannon多样性指数 Shannon diversity index
土样深度 Soil depth (cm)	NTS	SNTS	TT	NTS	SNTS	TT	NTS	SNTS	TT
0~20	7.3aA	7.7aA	5.3aB	111.7aAB	125.0aA	90.3aC	1.79aA	1.83aA	1.67aB
20~40	7.0aA	7.3aA	5.0aB	47.7bAB	57.3bA	43.7bC	1.75aA	1.77abA	1.65aA
40~60	3.7bB	4.7bA	3.0bC	22.3cAB	28.7cA	15.7cC	1.69aA	1.77abA	1.08bB
60~80	3.3bB	4.0bA	2.3cC	11.0dA	11.7dA	7.3dB	1.69aA	1.65bA	1.08bB
80~100	2.7cB	3.3cA	1.7dC	7.7eAB	12.0dA	4.3eB	0.77bB	1.0cA	0.55cC
100~120	1.0dA	1.0dA	0.7eB	4.7efB	11.0dA	1.7eB	0.77bB	1.0cA	0.54cC
120~140	0.7dA	1.0dA	0.0fA	2.3fgAB	4.0eA	0.0eB	0.0cA	0.0dA	0.0dA
140~160	0.0eA	0.7dA	0.0fA	0.0gA	3.0efA	0.0eA	0.0cA	0.0dA	0.0dA
160~180	0.0eA	0.0eA	0.0fA	0.0gA	0.0fA	0.0eA	0.0cA	0.0dA	0.0dA
180~200	0.0eA	0.0eAS	0.0fA	0.0gA	0.0fA	0.0eA	0.0cA	0.0dA	0.0dA
??NTS:免耕覆盖; SNTS:深松免耕覆盖; TT:传统耕作。同列不同小写字母表示不同土层间差异显著(P < 0.05);同行不同大写字母表示同一土层不同处理间差异显著(P < 0.05)。NTS: no-till with straw mulching; SNTS: no-till with straw mulching after subsoiling; TT: traditional tillage. Different small letters in the same column mean significant differences among different soil layers in the same treatment at P < 0.05, and different capital letters in the same row mean significant differences in the same soil layer among different treatments at P < 0.05.

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表4不同耕作措施下小麦不同生育期丛枝菌根真菌(AMF)侵染率和土壤孢子密度
Table4.Colonization rate and soil spore density of arbuscular mycorrhizal fungi (AMF) under different tillage treatment at different growth periods of wheat

生育期 Growing period	AMF侵染率Colonization rate of AMF (%)			孢子密度Spore density [spores?(100g)^-1(soil)]
生育期 Growing period	NTS	SNTS	TT	NTS	SNTS	TT
苗期Seedling	6.6bA	7.6bA	6.1bA	76.7cA	82.0dA	46.0cB
返青期Returning green	7.2bAB	8.6bA	6.3bB	70.7cA	80.7dA	43.7cB
拔节期Jointing	14.9aAB	16.1aA	10.6aB	81.3cAB	94.3cA	53.3cC
灌浆期Filling	13.8aAB	15.6aA	10.1aB	97.7bA	108.7bA	73.0bB
成熟期Maturity	0.0cA	0.0cA	0.0cA	111.7aAB	125.0aA	90.3aC
??NTS:免耕覆盖; SNTS:深松免耕覆盖; TT:传统耕作。同列不同小写字母表示不同土层间差异显著(P < 0.05);同行不同大写字母表示同一土层不同处理间差异显著(P < 0.05)。NTS: no-till with straw mulching; SNTS: no-till with straw mulching after subsoiling; TT: traditional tillage. Different small letters in the same column mean significant differences among different soil layers in the same treatment at P < 0.05, and different capital letters in the same row mean significant differences in the same soil layer among different treatments at P < 0.05.

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