不同生育时期水稻土壤细菌群落特征与生态功能预测

doi:10.7668/hbnxb.20193000

摘要/Abstract

摘要：

分蘖期和拔节—孕穗期是水稻产量构成要素中穗数和穗粒数形成的2个关键生育时期,氮素影响分蘖成穗和穗粒数的形成,细菌是土壤氮素循环的重要参与者。以高产水稻品种Y两优900和早丰优69为试验材料,通过设置2个氮肥施用比例(基蘖肥和穗粒肥比例分别为7:3和6:4),分析了分蘖期和拔节—孕穗期土壤细菌数量、群落特征的差异,及其与水稻产量、土壤氮素的关系。结果表明,水稻的分蘖期和拔节—孕穗期土壤细菌群落结构存在差异;分蘖期和拔节—孕穗期稻田土壤细菌优势菌门是变形菌门、拟杆菌门、绿弯菌门和酸杆菌门等。分蘖期绿弯菌门的相对丰度比拔节—孕穗期高6.16百分点,酸杆菌门的相对丰度比拔节—孕穗期高2.65百分点;拔节—孕穗期的变形菌门相对丰度比分蘖期高0.69百分点,拟杆菌门相对丰度比分蘖期高1.09百分点。相关性分析发现,水稻产量与分蘖期土壤细菌数量呈显著负相关关系,而与拔节—孕穗期土壤细菌数量和全氮含量均呈显著正相关关系;有效穗数与分蘖期土壤细菌数量呈显著正相关关系,穗粒数与分蘖期、拔节—孕穗期的氨氧化潜势均呈显著正相关关系。冗余分析(RDA)结果表明,稻田土壤细菌群落结构受多种因素共同影响,分蘖期土壤的铵态氮和拔节—孕穗期土壤的氨氧化潜势是显著影响稻田土壤细菌群落结构的2个环境因子。FAPROTAX功能预测进一步说明,分蘖期7:3处理的反硝化功能增强,其中Y两优900土壤细菌的反硝化功能最强。因此,分蘖期土壤细菌数量的增加可提高水稻有效穗数。拔节—孕穗期土壤细菌参与的氨氧化作用可促进水稻穗粒数的增加。高产水稻品种Y两优900和早丰优69在不同施氮比例下,产量、有效穗数和穗粒数的形成与土壤细菌数量、群落组成及其生态功能关系密切。

关键词: 稻田土壤, 16S rRNA扩增子测序, 细菌群落结构, 产量, 产量构成要素, 生育时期

Abstract:

Tillering stage and jointing—booting stage are important for the formation of panicle number and spikelets per panicle of rice,respectively.Nitrogen application affects the formation of panicle number and spikelet per panicle.Bacteria involve in soil nitrogen cycle.Two high-yield rice varieties,Y Liangyou 900 and Zaofengyou 69,were used to study the difference of soil bacterial number and community structure between tillering stage and jointing—booting stage,and the relationship between them and soil nitrogen,rice yield under two application ratios of nitrogen fertilizer(7:3 and 6:4 application ratios of basic tiller fertilizer to panicle-spikelet fertilizer).The results showed that there were significant differences in the structures of dominant bacterial groups in paddy soil between the tillering stage and jointing—booting stage,which were mainly Proteobacteria,Bacteroidetes,Chloroflexi and Acidobacteria.The relative abundance of Chloroflexi at the tillering stage was 6.16 percent points higher than that at the jointing—booting stage.The relative abundance of Acidobacteria at the tillering stage was 2.65 percent points higher than that at the jointing—booting stage.The relative abundance of Proteobacteria at the jointing—booting stage was 0.69 percent points higher than that at the tilling stage.The relative abundance of Bacteroidetes at the jointing—booting stage was 1.09 percent points higher than that at the tillering stage.The results of correlation analysis showed that rice yield was significantly negatively correlated with the number of soil bacteria at the tillering stage,but significantly positively correlated with the number of bacteria and total nitrogen content in the soil at the jointing—booting stage.There was a significant positive correlation between the number of effective panicles and the number of soil bacteria at the tillering stage.The number of spikelets per panicle was significantly positively correlated with the ammonia oxidation potential at the tillering stage and the jointing—booting stage.Redundancy analysis(RDA)showed that the bacterial community structures were affected by many factors.The contents of soil ammonium nitrogen and the ammonia oxidation potential were the main factors affecting the bacterial community structures at the tillering stage and the jointing—booting stage,respectively.The function prediction of FAPROTAX further indicated that the denitrification in the 7:3 treatment at the tillering stage was enhanced,especially in the soil of Y Liangyou 900.Therefore,increasing the number of soil bacteria at the tillering stage could enhance the number of effective panicles.Ammonia oxidation involved by soil bacteria at the jointing—booting stage can promote the increase of spikelets per panicle in rice.For the high-yielding rice varieties,Y Liangyou 900 and Zaofengyou 69 under different nitrogen application ratios,the formation of yield,effective panicle number and spikelets per panicle is closely related to soil bacterial number,community composition and ecological function.

Key words: Paddy soil, 16S rRNA amplicon sequencing, Bacterial community structure, Yield, Yield components, Growth stage

陈秋圆, 赵帅兵, 梁贺, 刘晔, 崔燕妮, 彭廷, 赵全志, 张静. 不同生育时期水稻土壤细菌群落特征与生态功能预测[J]. 华北农学报, 2022, 37(5): 140-149. doi: 10.7668/hbnxb.20193000.

CHEN Qiuyuan, ZHAO Shuaibing, LIANG He, LIU Ye, CUI Yanni, PENG Ting, ZHAO Quanzhi, ZHANG Jing. Soil Bacterial Community Characteristics and Ecological Function Prediction at Different Rice Growth Stages[J]. Acta Agriculturae Boreali-Sinica, 2022, 37(5): 140-149. doi: 10.7668/hbnxb.20193000.

http://www.hbnxb.net/CN/Y2022/V37/I5/140

图/表 9

表1 不同处理土壤生化因子含量变化

Tab.1 Changes in the content of soil biochemical factors in different treatments

时期 Stage	水稻品种 Rice variety	氮肥处理 N treatment	氨氧化潜势/ (mg/kg) AOP	硝化强度/ (mg/kg) NC	硝态氮/ (mg/kg) N $O 3 -$ -N	铵态氮/ (mg/kg) N $H 4 +$ -N	全氮/ (g/kg) TN	细菌数量/ (×10⁵ cfu/g) No. of bacteria
分蘖期	Y两优900	6:4	1.09±0.24b	6.21±0.22c	3.45±0.71a	20.22±3.60a	1.38±0.39ab	78.32±9.68b
Tillering stage		7:3	1.34±0.06b	6.59±0.44bc	2.33±1.26a	15.07±2.52b	1.72±0.06a	103.33±4.85a
	早丰优69	6:4	1.92±0.15b	7.91±0.87ab	4.19±1.34a	19.66±0.52a	1.15±0.31b	70.20±7.62b
		7:3	4.00±1.62a	9.48±1.35a	2.78±1.42a	14.99±1.54b	1.42±0.13ab	80.84±2.69b
拔节—孕穗期	Y两优900	6:4	1.20±0.05b	7.78±0.23a	1.64±0.58ab	8.88±0.67c	0.69±0.23bc	74.12±8.93c
Jointing—booting		7:3	1.12±0.07b	7.76±0.22a	1.31±0.26b	8.80±0.48c	0.65±0.25c	79.68±1.85bc
stage	早丰优69	6:4	1.58±0.22a	5.58±0.71b	1.64±0.67ab	12.28±0.24a	1.07±0.19ab	87.17±2.18b
		7:3	1.71±0.08a	5.93±1.17b	2.57±0.31a	10.36±0.66b	1.17±0.18a	117.95±5.09a
F值 F value	时期		8.11^*	6.19^*	13.67^**	114.07^**	29.01^**	6.96^*
	品种		21.58^**	0.21	2.65	2.43	0.95	4.34
	氮肥		6.11^*	3.23	1.67	18.17^**	3.06	52.47^**
	时期×氮肥		5.67^*	1.65	4.30	7.96^*	2.00	0.01
	时期×品种		6.78^*	46.52^**	0.02	4.06	14.06^**	67.96^**
	氮肥×品种		4.48^*	1.52	0.41	0.24	0.03	1.19
	时期×氮肥×品种		2.82	0.42	1.04	0.70	0.26	15.85^**

表1 不同处理土壤生化因子含量变化

Tab.1 Changes in the content of soil biochemical factors in different treatments

时期 Stage	水稻品种 Rice variety	氮肥处理 N treatment	氨氧化潜势/ (mg/kg) AOP	硝化强度/ (mg/kg) NC	硝态氮/ (mg/kg) N $O 3 -$ -N	铵态氮/ (mg/kg) N $H 4 +$ -N	全氮/ (g/kg) TN	细菌数量/ (×10⁵ cfu/g) No. of bacteria
分蘖期	Y两优900	6:4	1.09±0.24b	6.21±0.22c	3.45±0.71a	20.22±3.60a	1.38±0.39ab	78.32±9.68b
Tillering stage		7:3	1.34±0.06b	6.59±0.44bc	2.33±1.26a	15.07±2.52b	1.72±0.06a	103.33±4.85a
	早丰优69	6:4	1.92±0.15b	7.91±0.87ab	4.19±1.34a	19.66±0.52a	1.15±0.31b	70.20±7.62b
		7:3	4.00±1.62a	9.48±1.35a	2.78±1.42a	14.99±1.54b	1.42±0.13ab	80.84±2.69b
拔节—孕穗期	Y两优900	6:4	1.20±0.05b	7.78±0.23a	1.64±0.58ab	8.88±0.67c	0.69±0.23bc	74.12±8.93c
Jointing—booting		7:3	1.12±0.07b	7.76±0.22a	1.31±0.26b	8.80±0.48c	0.65±0.25c	79.68±1.85bc
stage	早丰优69	6:4	1.58±0.22a	5.58±0.71b	1.64±0.67ab	12.28±0.24a	1.07±0.19ab	87.17±2.18b
		7:3	1.71±0.08a	5.93±1.17b	2.57±0.31a	10.36±0.66b	1.17±0.18a	117.95±5.09a
F值 F value	时期		8.11^*	6.19^*	13.67^**	114.07^**	29.01^**	6.96^*
	品种		21.58^**	0.21	2.65	2.43	0.95	4.34
	氮肥		6.11^*	3.23	1.67	18.17^**	3.06	52.47^**
	时期×氮肥		5.67^*	1.65	4.30	7.96^*	2.00	0.01
	时期×品种		6.78^*	46.52^**	0.02	4.06	14.06^**	67.96^**
	氮肥×品种		4.48^*	1.52	0.41	0.24	0.03	1.19
	时期×氮肥×品种		2.82	0.42	1.04	0.70	0.26	15.85^**

表2 不同处理水稻产量及其构成要素

Tab.2 Rice yield and yield components in different treatments

水稻品种 Rice variety	氮肥处理 N treatment	有效穗数/ (万穗/hm²) No. of effective panicles	穗粒数 Spikelets per panicle	结实率/% Seed-setting rate	千粒质量/g 1000-grain weight	产量/ (t/hm²) Yield
Y两优900	6:4	212.92±13.50a	188.53±20.36c	75.13±2.66b	23.13±0.47a	10.98±1.23ab
Y Liangyou 900	7:3	229.98±6.43a	185.55±10.93c	74.69±3.64b	22.90±0.12a	10.81±0.66b
早丰优69	6:4	225.17±25.03a	221.05±8.22b	83.31±1.88a	22.68±0.11a	11.80±0.53ab
Zaofengyou 69	7:3	234.00±23.03a	248.84±22.54a	83.67±1.94a	22.77±0.47a	11.96±0.26a
F值 F value	品种	0.08	33.16^**	53.29^**	3.65	7.41^*
	氮肥	2.99	0.74	0.01	0.20	0.61
	品种×氮肥	1.52	1.68	0.12	1.07	0.67

表3 土壤生化因子与产量及其构成要素的相关性分析(n=12)

Tab.3 Correlation analysis between soil biochemistry indexes and yield,its components

指标 Index	生育时期 Stage	有效穗数 No. of effective panicles	穗粒数 Spikelets per panicle	产量 Yield
细菌数量	分蘖期	0.61^*	-0.41	-0.65^*
No. of bacteria	拔节—孕穗期	0.14	0.52	0.62^*
氨氧化潜势	分蘖期	0.16	0.64^*	0.36
AOP	拔节—孕穗期	-0.10	0.65^*	0.32
硝化强度	分蘖期	0.26	0.57	0.49
NC	拔节—孕穗期	-0.02	-0.35	-0.53
硝态氮	分蘖期	-0.40	0.03	0.04
N $O 3 -$ -N	拔节—孕穗期	-0.29	0.47	0.22
铵态氮	分蘖期	-0.20	0.01	-0.07
N $H 4 +$ -N	拔节—孕穗期	0.11	0.53	0.28
全氮	分蘖期	0.06	-0.69^*	-0.10
TN	拔节—孕穗期	-0.02	0.43	0.64^*

表3 土壤生化因子与产量及其构成要素的相关性分析(n=12)

Tab.3 Correlation analysis between soil biochemistry indexes and yield,its components

指标 Index	生育时期 Stage	有效穗数 No. of effective panicles	穗粒数 Spikelets per panicle	产量 Yield
细菌数量	分蘖期	0.61^*	-0.41	-0.65^*
No. of bacteria	拔节—孕穗期	0.14	0.52	0.62^*
氨氧化潜势	分蘖期	0.16	0.64^*	0.36
AOP	拔节—孕穗期	-0.10	0.65^*	0.32
硝化强度	分蘖期	0.26	0.57	0.49
NC	拔节—孕穗期	-0.02	-0.35	-0.53
硝态氮	分蘖期	-0.40	0.03	0.04
N $O 3 -$ -N	拔节—孕穗期	-0.29	0.47	0.22
铵态氮	分蘖期	-0.20	0.01	-0.07
N $H 4 +$ -N	拔节—孕穗期	0.11	0.53	0.28
全氮	分蘖期	0.06	-0.69^*	-0.10
TN	拔节—孕穗期	-0.02	0.43	0.64^*

图1 不同处理稻田土壤细菌群落结构的PCoA分析 Y.Y两优900;Z.早丰优69;T.分蘖期;J.拔节—孕穗期。图2—5同。

Fig.1 Principal coordinate analysis of bacterial community structures in paddy soils of different treatments Y.Y Liangyou 900;Z.Zaofengyou 69;T.Tillering stage; J.Jointing—booting stage.The same as Fig.2—5.

图2 不同处理稻田土壤细菌优势菌门/纲分析 Y-6:4.氮肥比例6:4处理的Y两优900;Z-6:4.氮肥比例6:4处理的早丰优69;Y-7:3.氮肥比例7:3处理的Y两优900;Z-7:3.氮肥比例7:3处理的早丰优69。其他指相对丰度排名前10以外的其他细菌。因变形菌门丰度较高,相对丰度为纲水平,其余为门水平。

Fig.2 Relative abundance of dominant bacteria at phyla/class level in paddy soils of different treatments Y-6:4.Y Liangyou 900 treated with nitrogen fertilizer ratio of 6:4;Z-6:4.Zaofengyou 69 treated with nitrogen fertilizer ratio of 6:4;Y-7:3.Y Liangyou 900 treated with nitrogen fertilizer ratio of 7:3;Z-7:3.Zaofengyou 69 treated with nitrogen fertilizer ratio of 7:3.Others represents the rest of soil bacteria except the top 10 bacteria in the relative abundance.Data of Proteobacteria are used at class level due to its high abundance,and the rests are used at phylum level.

图3 不同处理差异ASV及对应门水平分类组成 A.2个时期Y两优900土壤富集的ASV;B.2个时期Y两优900土壤富集ASV的组成;C.2个时期早丰优69土壤富集的ASV;D.2个时期早丰优69土壤富集ASV的组成;E.拔节—孕穗期土壤富集的ASV;F.拔节—孕穗期土壤富集ASV的组成。G.分蘖期土壤富集的ASV;H.分蘖期土壤富集ASV的组成。

Fig.3 The overlap and taxonomic composition of ASVs enriched at phylum level under different treatments A.ASVs enriched in the soil of Y Liangyou 900 in the two periods;B.Composition of ASVs enriched in the soil of Y Liangyou 900 in the two periods;C.ASVs enriched in the soil of Zaofengyou 69 in the two stages;D.Composition of ASVs enriched in the soil of Zaofengyou 69 in the two stages;E.ASVs enriched in the soil at the jointing—booting stage;F.Composition of ASVs enriched in the soil at the jointing—booting stage;G.ASVs enriched in the soil at the tillering stage;H.Composition of ASVs enriched in the soil at the tillering stage.

图4 不同处理环境因子与稻田土壤优势细菌群落的冗余分析 A.分蘖期环境因子与稻田土壤细菌群落结构的RDA分析;B.拔节—孕穗期环境因子与稻田土壤细菌群落结构的RDA分析;C.Y两优900品种环境因子与稻田土壤细菌群落结构的RDA分析;D.早丰优69品种环境因子与稻田土壤细菌群落结构的RDA分析。

Fig.4 Redundancy analysis of environmental factors and dominant bacterial communities in paddy soils of different treatments A.RDA analysis of environmental factors and community structure of soil bacteria at tillering stage;B.RDA analysis of environmental factors and community structure of soil bacteria at jointing—booting stage;C.RDA analysis of environmental factors and community structure of soil bacteria of Y Liangyou 900;D.RDA analysis of environmental factors and community structure of soil bacteria of Zaofengyou 69.

表4 不同处理环境因子对稻田土壤优势细菌群落的解释率

Tab.4 Explanation rate of environmental factors on the dominant bacterial communities in paddy soils of different treatments%

处理 Treatment	氨氧化潜势 AOP	硝化强度 NC	硝态氮 N $O 3 -$ -N	铵态氮 N $H 4 +$ -N	全氮 TN
分蘖期Tillering stage	15.88	22.01	12.19	35.64^**	14.28
拔节—孕穗期Jointing—booting stage	29.57^*	18.75	19.79	15.92	15.97
Y两优900 Y Liangyou 900	2.17	54.33	16.23	9.75	17.52
早丰优69 Zaofengyou 69	4.76	25.62	9.17	45.69	14.76

表4 不同处理环境因子对稻田土壤优势细菌群落的解释率

Tab.4 Explanation rate of environmental factors on the dominant bacterial communities in paddy soils of different treatments%

处理 Treatment	氨氧化潜势 AOP	硝化强度 NC	硝态氮 N $O 3 -$ -N	铵态氮 N $H 4 +$ -N	全氮 TN
分蘖期Tillering stage	15.88	22.01	12.19	35.64^**	14.28
拔节—孕穗期Jointing—booting stage	29.57^*	18.75	19.79	15.92	15.97
Y两优900 Y Liangyou 900	2.17	54.33	16.23	9.75	17.52
早丰优69 Zaofengyou 69	4.76	25.62	9.17	45.69	14.76

图5 不同处理间稻田土壤细菌功能相对丰度差异热图

Fig.5 Heat map of differences in relative abundances of soil bacterial functions in paddy soils among different treatments

参考文献 54

[1]	Yang J C, Zhang H, Zhang J H. Root morphology and physiology in relation to the yield formation of rice[J]. Journal of Integrative Agriculture, 2012(6):920-926.doi: 10.1016/S2095-3119(12)60082-3. doi: 10.1016/S2095-3119(12)60082-3
[2]	崔月贞, 吴玉红, 郝兴顺, 张春辉, 秦宇航, 王晓娥, 杨云霞. 不同有机氮替代部分无机氮对水稻产量及土壤微生物的影响[J]. 西北农业学报, 2019, 28(10):1689-1697.doi: 10.7606/j.issn.1004-1389.2019.10.016. doi: 10.7606/j.issn.1004-1389.2019.10.016
	Cui Y Z, Wu Y H, Hao X S, Zhang C H, Qin Y H, Wang X E, Yang Y X. Effects of partial replacement of chemical fertilizer by organic manure on rice yield and soil microorganisms in Hanzhong Basin[J]. Acta Agriculturae Boreali-Occidentalis Sinica, 2019, 28(10):1689-1697.
[3]	胡桂萍. 水稻内生菌及其根系土壤微生物群落多样性的研究[D]. 福州: 福建农林大学, 2010.
	Hu G P. Study on the community diversity of the endophytic and rhizosphere microorganism of rice (Oryza sativa L.)[D]. Fuzhou: Fujian Agriculture and Forestry University, 2010.
[4]	雍太文, 杨文钰, 向达兵, 陈小容. 不同种植模式对作物根系生长、产量及根际土壤微生物数量的影响[J]. 应用生态学报, 2012, 23(1):125-132.doi: 10.13287/j.1001-9332.2012.0017. doi: 10.13287/j.1001-9332.2012.0017
	Yong T W, Yang W Y, Xiang D B, Chen X R. Effects of different cropping modes on crop root growth,yield,and rhizosphere soil microbes' number[J]. Chinese Journal of Applied Ecology, 2012, 23(1):125-132.
[5]	陈龙怀. 氮肥运筹调控下水稻根际土壤生物学特性分析[D]. 福州: 福建农林大学, 2012.
	Chen L H. The microbial characteristics analysis of rice rhizosphere soil under nitrogen regulation[D]. Fuzhou: Fujian Agriculture and Forestry University, 2012.
[6]	Edwards J, Johnson C, Santos-Medellín C, Lurie E, Podishetty N K, Bhatnagar S, Eisen J A, Sundaresan V. Structure,variation,and assembly of the root-associated microbiomes of rice[J]. Proceedings of the National Academy of Sciences of the United States of America, 2015, 112(8):E911-E920.doi: 10.1073/pnas.1414592112. doi: 10.1073/pnas.1414592112
[7]	Zhang J Y, Liu Y X, Zhang N, Hu B, Jin T, Xu H R, Qin Y, Yan P X, Zhang X N, Guo X X, Hui J, Cao S Y, Wang X, Wang C, Wang H, Qu B Y, Fan G Y, Yuan L X, Garrido-Oter R, Chu C C, Bai Y. NRT1.1B is associated with root microbiota composition and nitrogen use in field-grown rice[J]. Nature Biotechnology, 2019, 37(6):676-684.doi: 10.1038/s41587-019-0104-4. doi: 10.1038/s41587-019-0104-4 pmid: 31036930
[8]	Edwards J A, Santos-Medellín C M, Liechty Z S, Nguyen B, Lurie E, Eason S, Phillips G, Sundaresan V. Compositional shifts in root-associated bacterial and archaeal microbiota track the plant life cycle in field-grown rice[J]. PLoS Biology, 2018, 16(2):e2003862.doi: 10.1371/journal.pbio.2003862. doi: 10.1371/journal.pbio.2003862 URL
[9]	Chen S M, Waghmode T R, Sun R B, Kuramae E E, Hu C S, Liu B B. Root-associated microbiomes of wheat under the combined effect of plant development and nitrogen fertilization[J]. Microbiome, 2019, 7(1):136.doi: 10.1186/s40168-019-0750-2. doi: 10.1186/s40168-019-0750-2 pmid: 31640813
[10]	Shi Y, Li Y T, Xiang X J, Sun R B, Yang T, He D, Zhang K P, Ni Y Y, Zhu Y G, Adams J M, Chu H Y. Spatial scale affects the relative role of stochasticity versus determinism in soil bacterial communities in wheat fields across the North China Plain[J]. Microbiome, 2018, 6(1):27.doi: 10.1186/s40168-018-0409-4. doi: 10.1186/s40168-018-0409-4 pmid: 29402331
[11]	Sun R B, Li W Y, Dong W X, Tian Y P, Hu C S, Liu B B. Tillage changes vertical distribution of soil bacterial and fungal communities[J]. Frontiers in Microbiology, 2018, 9:699.doi: 10.3389/fmicb.2018.00699. doi: 10.3389/fmicb.2018.00699 pmid: 29686662
[12]	梁继华, 黄白红, 田学辉. 不同土壤改良剂对玉米产量及根际土壤微生物和酶活性的影响[J]. 山东农业科学, 2022, 54(1):80-85.doi: 10.14083/j.issn.1001-4942.2022.01.013. doi: 10.14083/j.issn.1001-4942.2022.01.013
	Liang J H, Huang B H, Tian X H. Effects of different soil conditioners on maize yield and microorganism and enzyme activity in rhizosphere soil[J]. Shandong Agricultural Sciences, 2022, 54(1):80-85.
[13]	Müller D B, Vogel C, Bai Y, Vorholt J A. The plant microbiota:Systems-level insights and perspectives[J]. Annual Review of Genetics, 2016, 50:211-234.doi: 10.1146/annurev-genet-120215-034952. doi: 10.1146/annurev-genet-120215-034952 URL
[14]	Liu Y X, Qin Y, Bai Y. Reductionist synthetic community approaches in root microbiome research[J]. Current Opinion in Microbiology, 2019, 49:97-102.doi: 10.1016/j.mib.2019.10.010. doi: 10.1016/j.mib.2019.10.010 URL
[15]	Yuan J, Zhao J, Wen T, Zhao M L, Li R, Goossens P, Huang Q W, Bai Y, Vivanco J M, Kowalchuk G A, Berendsen R L, Shen Q R. Root exudates drive the soil-borne legacy of aboveground pathogen infection[J]. Microbiome, 2018, 6(1):156.doi: 10.1186/s40168-018-0537-x. doi: 10.1186/s40168-018-0537-x pmid: 30208962
[16]	胡雅丽, 戴睿, 刘永鑫, 张婧赢, 胡斌, 储成才, 袁怀波, 白洋. 水稻典型品种日本晴和IR24根系微生物组的解析[J]. 遗传, 2020, 42(5):506-518.doi: 10.16288/j.yczz.20-070. doi: 10.16288/j.yczz.20-070
	Hu Y L, Dai R, Liu Y X, Zhang J Y, Hu B, Chu C C, Yuan H B, Bai Y. Analysis of rice root bacterial microbiota of Nipponbare and IR24[J]. Hereditas, 2020, 42(5):506-518.
[17]	寿南松, 黄迪, 吴漪, 陈若夫, 张立钦, 吴小映. 不同施氮水平下配施硅肥对水稻根部周围土壤微生物群落结构的影响[J]. 土壤通报, 2021, 52(4):903-911.doi: 10.19336/j.cnki.trtb.2020101001. doi: 10.19336/j.cnki.trtb.2020101001
	Shou N S, Huang D, Wu Y, Chen R F, Zhang L Q, Wu X Y. Effects of silicon on microbial community structure of rhizosphere soil of rice at different nitrogen levels[J]. Chinese Journal of Soil Science, 2021, 52(4):903-911.
[18]	袁红朝, 吴昊, 葛体达, 李科林, 吴金水, 王久荣. 长期施肥对稻田土壤细菌、古菌多样性和群落结构的影响[J]. 应用生态学报, 2015, 26(6):1807-1813.doi: 10.13287/j.1001-9332.20150413.008. doi: 10.13287/j.1001-9332.20150413.008
	Yuan H Z, Wu H, Ge T D, Li K L, Wu J S, Wang J R. Effects of long-term fertilization on bacterial and archaeal diversity and community structure within subtropical red paddy soils[J]. Chinese Journal of Applied Ecology, 2015, 26(6):1807-1813.
[19]	潘丽媛, 肖炜, 董艳, 李龚程, 张乃明, 段红平, 张仕颖. 超高产生态区水稻根际微生物物种及功能多样性研究[J]. 农业资源与环境学报, 2016, 33(6):583-590.doi: 10.13254/j.jare.2016.0074. doi: 10.13254/j.jare.2016.0074
	Pan L Y, Xiao W, Dong Y, Li G C, Zhang N M, Duan H P, Zhang S Y. Microbial species and functional diversity in rice rhizosphere of high-yield special ecological areas[J]. Journal of Agricultural Resources and Environment, 2016, 33(6):583-590.
[20]	马微微, 陈灿, 黄璜, 任勃, 周晶, 李桂香, 王忍. 垄作稻鱼鸡共生对稻田土壤养分含量及水稻产量的影响[J]. 河南农业科学, 2021, 50(8):9-17.doi: 10.15933/j.cnki.1004-3268.2021.08.002. doi: 10.15933/j.cnki.1004-3268.2021.08.002
	Ma W W, Chen C, Huang H, Ren B, Zhou J, Li G X, Wang R. Effect of rice-fish-chicken symbiosis on paddy soil nutrient and rice yield under ridge cultivation[J]. Journal of Henan Agricultural Sciences, 2021, 50(8):9-17.
[21]	蒋鹏, 徐富贤, 张林, 周兴兵, 朱永川, 郭晓艺, 刘茂, 陈琳, 张容, 熊洪. 高温高湿区增密减氮对杂交稻内6优107产量形成和氮肥利用率的影响[J]. 中国生态农业学报(中英文), 2021, 29(10):1679-1691.doi: 10.13930/j.cnki.cjea.210288. doi: 10.13930/j.cnki.cjea.210288
	Jiang P, Xu F X, Zhang L, Zhou X B, Zhu Y C, Guo X Y, Liu M, Chen L, Zhang R, Xiong H. Effect of increased plant density with reduced nitrogen on yield formation and nitrogen use efficiency of hybrid rice under high temperature and high humidity conditions[J]. Chinese Journal of Eco-Agriculture, 2021, 29(10):1679-1691.
[22]	Wang Q X, Li M L, Chen X J, Su Y, Yu M, Shen A L. Effects of nitrogen management on soil microbial community structure at different growth stages under straw returning in paddy soils[J]. Ying Yong Sheng Tai Xue Bao, 2020, 31(3):935-944.doi: 10.13287/j.1001-9332.202003.027. doi: 10.13287/j.1001-9332.202003.027
[23]	中国科学院南京土壤研究所. 土壤理化分析[M]. 上海: 上海科学技术出版社, 1978.
	Nanjing Institute of Soil Science, Chinese Academy of Sciences. Soil physical and chemical analysis[M]. Shanghai: Science and Technology Press, 1978.
[24]	国家环保局本书编委会. 水和废水监测分析方法[M]. 北京: 中国环境科学出版社, 1989.
	National Environmental Protection Agency Book Edition Committee. Water and wastewater monitoring and analysis methods[M]. Beijing: China Environmental Science Press, 1989.
[25]	Li Y Y, Chapman S J, Nicol G W, Yao H Y. Nitrification and nitrifiers in acidic soils[J]. Soil Biology and Biochemistry, 2018, 116:290-301.doi: 10.1016/j.soilbio.2017.10.023. doi: 10.1016/j.soilbio.2017.10.023 URL
[26]	许光辉, 郑洪元. 土壤微生物分析方法手册[M]. 北京: 农业出版社, 1986.
	Xu G H, Zheng H Y. Handbook of soil microbial analysis methods[M]. Beijing: Agricultural Press, 1986.
[27]	中国科学院南京土壤研究所微生物室. 土壤微生物研究法[M]. 北京: 科学出版社, 1985.
	Microbiology Laboratory,Nanjing Institute of Soil Science,Chinese Academy of Sciences. Soil microbiology research method[M]. Beijing: Science Press, 1985.
[28]	Eldridge D J, Travers S K, Val J, Ding J Y, Wang J T, Singh B K, Delgado-Baquerizo M. Experimental evidence of strong relationships between soil microbial communities and plant germination[J]. Journal of Ecology, 2021, 109(6):2488-2498.doi: 10.1111/1365-2745.13660. doi: 10.1111/1365-2745.13660 URL
[29]	伍泳彰, 陈杰. 一种用于构建16SrRNA基因扩增子测序文库的引物组及构建方法[P].CN107829146B, 2021-07-09.
	Wu Y Z, Chen J. A primer set and construction method for constructing 16SrRNA gene amplicon sequencing library[P].CN107829146B, 2021-07-09.
[30]	Fadrosh D W, Ma B, Gajer P, Sengamalay N, Ott S, Brotman R M, Ravel J. An improved dual-indexing approach for multiplexed 16S rRNA gene sequencing on the Illumina MiSeq platform[J]. Scientific Reports, 2014, 2(1):6.doi: 10.1186/2049-2618-2-6. doi: 10.1186/2049-2618-2-6
[31]	Bolyen E, Rideout J R, Dillon M R, Bokulich N A, Abnet C C, Al-Ghalith G A, et al. Reproducible,interactive,scalable and extensible microbiome data science using QIIME 2[J]. Nat Biotechnol, 2019, 37(8):852-857.doi: 10.1038/s41587-019-0209-9. doi: 10.1038/s41587-019-0209-9 pmid: 31341288
[32]	Dixon P. vegan,a package of R functions for community ecology[J]. Journal of Vegetation Science, 2003, 14(6):927-930.doi: 10.1111/j.1654-1103.2003.tb02228.x. doi: 10.1111/j.1654-1103.2003.tb02228.x URL
[33]	Robinson M D, McCarthy D J, Smyth G K. edgeR:a Bioconductor package for differential expression analysis of digital gene expression data[J]. Bioinformatics, 2010, 26(1):139-140.doi: 10.1093/bioinformatics/btp616. doi: 10.1093/bioinformatics/btp616 pmid: 19910308
[34]	Gómez-Rubio V. ggplot2-elegant graphics for data analysis[J]. Journal of Statistical Software, 2017, 77:1-3.doi: 10.1111/J.1467-985X.2010.00676_9.X. doi: 10.1111/J.1467-985X.2010.00676_9.X
[35]	Louca S, Parfrey L W, Doebeli M. Decoupling function and taxonomy in the global ocean microbiome[J]. Science, 2016, 353(6305):1272-1277.doi: 10.1126/science.aaf4507. doi: 10.1126/science.aaf4507 pmid: 27634532
[36]	Galili T, O'Callaghan A, Sidi J, Sievert C. Heatmaply:an R package for creating interactive cluster heatmaps for online publishing[J]. Bioinformatics, 2017, 34(9):1600-1602.doi: 10.1093/bioinformatics/btx657. doi: 10.1093/bioinformatics/btx657 URL
[37]	Singh B, Katyal J C, Malhotra P K, Vlek P L G. Path coefficient analysis of N nutrition on yield and yield components for rice in a highly percolating soil[J]. Communications in Soil Science and Plant Analysis, 1986, 17(8):853-867.doi: 10.1080/00103628609367757. doi: 10.1080/00103628609367757 URL
[38]	林忠秀, 易镇邪. 施氮量对优质晚稻盛泰优018产量形成与氮利用效率的影响[J]. 作物研究, 2021, 35(4):293-296.doi: 10.16848/j.cnki.issn.1001-5280.2021.04.02. doi: 10.16848/j.cnki.issn.1001-5280.2021.04.02
	Lin Z X, Yi Z X. Effect of nitrogen application rate on yield formation and nitrogen use efficiency of the high-quality late rice variety Shengtaiyou 018[J]. Crop Research, 2021, 35(4):293-296.
[39]	王磊, 朱德峰, 陈希春, 谢芙贤, 蔡体常, 高美黎, 纪希平, 张申龙, 郑朝华, 张狄标. 水稻产量和穗粒结构关系的估计[J]. 中国水稻科学, 1995, 9(1):33-38.doi: 10.16819/j.1001-7216.1995.01.006. doi: 10.16819/j.1001-7216.1995.01.006
	Wang L, Zhu D F, Chen X C, Xie F X, Cai T C, Gao M L, Ji X P, Zhang S L, Zheng C H, Zhang D B. Estimation of relationship between rice yield and panicle-grain structure[J]. Chinese Journal of Rice Science, 1995, 9(1):33-38.
[40]	王祎, 杨文浩, 毛艳玲, 周碧青, 聂三安, 邢世和. 水稻生育期对不同施肥条件下黄泥田土壤无机氮及细菌群落的影响[J]. 应用与环境生物学报, 2019, 25(6):1352-1358.doi: 10.19675/j.cnki.1006-687x.2019.01046. doi: 10.19675/j.cnki.1006-687x.2019.01046
	Wang Y, Yang W H, Mao Y L, Zhou B Q, Nie S N, Xing S H. Effect of the phenological stage of rice growth on soil-soluble inorganic nitrogen and bacterial communities in a yellow clayey soil under different fertilization patterns[J]. Chinese Journal of Applied and Environmental Biology, 2019, 25(6):1352-1358.
[41]	聂江文, 王幼娟, 吴邦魁, 刘章勇, 朱波. 双季稻田冬闲期土壤细菌数量及结构对施氮的响应[J]. 植物营养与肥料学报, 2019, 25(6):889-896.doi: 10.11674/zwyf.18229. doi: 10.11674/zwyf.18229
	Nie J W, Wang Y J, Wu B K, Liu Z Y, Zhu B. Response of soil bacterial quantity and community to nitrogen fertilization in double-cropping rice field during winter fallow period[J]. Journal of Plant Nutrition and Fertilizers, 2019, 25(6):889-896.
[42]	慕君, 马旭东, 张丹丹, 王敏鸽, 张霞, 张育林, 王旭东. 有机肥与化肥配施下土壤氮组分变化与氮素利用率研究[J]. 干旱地区农业研究, 2021, 39(5):107-113.doi: 10.7606/j.issn.1000-7601.2021.05.14. doi: 10.7606/j.issn.1000-7601.2021.05.14
	Mu J, Ma X D, Zhang D D, Wang M G, Zhang X, Zhang Y L, Wang X D. Study on nitrogen composition variation and nitrogen use efficiency under combined application of organic fertilizer and chemical fertilizer[J]. Agricultural Research in the Arid Areas, 2021, 39(5):107-113.
[43]	侯海军, 张文钊, 沈建林, 王聪, 秦红灵. 水分管理对稻田细菌丰度与群落结构的影响[J]. 生态环境学报, 2016, 25(9):1431-1438.doi: 10.16258/j.cnki.1674-5906.2016.09.002. doi: 10.16258/j.cnki.1674-5906.2016.09.002
	Hou H J, Zhang W Z, Shen J L, Wang C, Qin H L. Effect of water management on soil bacterial abundance and community in the rice paddy field[J]. Ecology and Environmental Sciences, 2016, 25(9):1431-1438.
[44]	吴晶. 氮肥施用量及种类对高产稻田土壤理化性质和细菌群落的影响[D]. 扬州: 扬州大学, 2019.
	Wu J. Effects of nitrogen application rates and types on soil physical and chemical properties and bacterial communities in high-yielding paddy fields[D]. Yangzhou: Yangzhou University, 2019.
[45]	朱国洁, 张娜, 杜雯, 李晓鹤, 王晨, 王欣丽. 氨氧化微生物在氮循环中的生态功能及其影响因子[J]. 天津农业科学, 2015, 21(12):48-53.doi: 10.3969/j.issn.1006-6500.2015.12.011. doi: 10.3969/j.issn.1006-6500.2015.12.011
	Zhu G J, Zhang N, Du W, Li X H, Wang C, Wang X L. Ecological function of ammonia oxidizing microorganisms in the nitrogen cycle and their influence factors[J]. Tianjin Agricultural Sciences, 2015, 21(12):48-53.
[46]	王丽娜. 转基因水稻根际固氮微生物群落结构分析[D]. 哈尔滨: 东北农业大学, 2009.
	Wang L N. Analysis of diazotroph communities in transgenic rice rhizosphere[D]. Harbin: Northeast Agricultural University, 2009.
[47]	栗笑阳, 郭夏丽. 不同施氮水平对小麦根际土壤氨氧化微生物的影响[J]. 河南农业科学, 2020, 49(12):69-76.doi: 10.15933/j.cnki.1004-3268.2020.12.010. doi: 10.15933/j.cnki.1004-3268.2020.12.010
	Li X Y, Guo X L. Effects of different nitrogen levels on ammonia oxidizing microorganisms in wheat rhizosphere soil[J]. Journal of Henan Agricultural Sciences, 2020, 49(12):69-76.
[48]	王智慧. 不同pH紫色土中硝化作用及硝化微生物宏基因组研究[D]. 重庆: 西南大学, 2017.
	Wang Z H. Metagenomic research of nitrifying microorganisms and nitrification in purple soils with different pH[D]. Chongqing: Southwest University, 2017.
[49]	Sun Y P, Guan Y T, Wang H Y, Wu G X. Autotrophic nitrogen removal in combined nitritation and Anammox systems through intermittent aeration and possible microbial interactions by quorum sensing analysis[J]. Bioresour Technol, 2019, 272:146-155.doi: 10.1016/j.biortech.2018.10.017. doi: 10.1016/j.biortech.2018.10.017 URL
[50]	Liu C Y, Zheng X H, Zhou Z X, Han S H, Wang Y H, Wang K, Liang W G, Li M, Chen D L, Yang Z P. Nitrous oxide and nitric oxide emissions from an irrigated cotton field in Northern China[J]. Plant and Soil, 2010, 332(1/2):123-134.doi: 10.1007/s11104-009-0278-5. doi: 10.1007/s11104-009-0278-5 URL
[51]	靳东升, 张强, 张变华, 郜春花, 李建华, 卢晋晶. 种植植物对煤矸石填埋区复垦土壤真菌多样性及养分含量的影响[J]. 华北农学报, 2020, 35(5):206-213.doi: 10.7668/hbnxb.20190997. doi: 10.7668/hbnxb.20190997
	Jin D S, Zhang Q, Zhang B H, Gao C H, Li J H, Lu J J. Effect of planting plants on fungi diversity and nutrients in reclaimed soil of coal gangue landfill area[J]. Acta Agriculturae Boreali-Sinica, 2020, 35(5):206-213. doi: 10.7668/hbnxb.20190997
[52]	张宸辉. 不同氮肥对油茶根际土壤氮循环微生物群落的影响[D]. 长沙: 中南林业科技大学, 2021.doi: 10.27662/d.cnki.gznlc.2021.000849. doi: 10.27662/d.cnki.gznlc.2021.000849
	Zhang C H. Effect of different nitrogen fertilizer on nitrogen circulation microrobial community of soil in Camellia oleifera[D]. Changsha: Central South University of Forestry & Technology, 2021.
[53]	姜红芳, 兰宇辰, 李猛, 赵洋, 李晓蕾, 王鹤璎, 徐令旗, 王君钰, 张佳柠, 吕艳东, 郭晓红. 氮肥运筹对盐碱地水稻籽粒氮代谢关键酶活性和蛋白质含量的影响[J]. 华北农学报, 2019, 34(S1):213-220.doi: 10.7668/hbnxb.20190719. doi: 10.7668/hbnxb.20190719
	Jiang H F, Lan Y C, Li M, Zhao Y, Li X L, Wang H Y, Xu L Q, Wang J Y, Zhang J N, Lü Y D, Guo X H. Effects of nitrogen fertilizer applications on key enzyme activities of nitrogen metabolism and protein content of rice in saline-alkali land[J]. Acta Agriculturae Boreali-Sinica, 2019, 34(S1):213-220. doi: 10.7668/hbnxb.20190719
[54]	宋亚娜, 林艳, 陈子强. 氮肥水平对稻田细菌群落及N₂O排放的影响[J]. 中国生态农业学报, 2017, 25(9):1266-1275.doi: 10.13930/j.cnki.cjea.170146. doi: 10.13930/j.cnki.cjea.170146
	Song Y N, Lin Y, Chen Z Q. Effect of nitrogen fertilizer level on bacterial community and N₂O emission in paddy soil[J]. Chinese Journal of Eco-Agriculture, 2017, 25(9):1266-1275.

选择文件类型/文献管理软件名称

选择包含的内容