保护播种下紫花苜蓿根际土壤氨氧化和反硝化微生物群落对糜子种植比例变化的响应

doi:10.11686/cyxb2024369

摘要/Abstract

摘要：

为研究保护播种下糜子种植比例对紫花苜蓿根际土壤因子、氨氧化微生物［氨氧化古菌（ammonia-oxidizing archaea，AOA）和氨氧化细菌（ammonia-oxidizing bacteria，AOB）］和反硝化微生物（nirK、nirS和nosZ）的影响，以黄土高原旱作农业区紫花苜蓿根际土壤为对象，设置4种糜子-紫花苜蓿种植比例［1∶1（1P1M），1∶2（1P2M），1∶3（1P3M）和2∶3（2P3M）］并以紫花苜蓿单播（M）为对照，利用高通量测序分析紫花苜蓿根际土壤中氨氧化和反硝化微生物群落多样性、结构、组成、共现网络及其与土壤因子的相关性。结果表明：随着糜子种植密度的增加，保护播种提高了土壤总氮含量和稳定碳氮同位素值，且在保护播种2P3M中显著增加（P<0.05），土壤有机碳含量先降低后增加，且在2P3M中显著下降（P<0.05）。保护播种提高了土壤氨氧化和nosZ微生物的丰富度，降低了nirK微生物的丰富度，其中，AOA微生物α多样性对种植比例变化更敏感。β多样性分析发现，不同保护播种比例之间，土壤氨氧化和反硝化微生物群落结构差异均不明显。AOA和AOB微生物分别以亚硝化球菌属和亚硝化螺菌属为优势属，其含量均超过85%；而反硝化微生物表现为富集促进植物生长发育和养分转化的菌属，如nirK微生物中无色杆菌属和nirS微生物中固氮弧菌属等。共现网络分析表明，保护播种2P3M处理的土壤氨氧化和反硝化微生物群落具有更复杂的共现网络，主要体现在网络复杂性和模块化系数指标。相关性分析进一步显示，全氮、钙离子、镁离子与AOA微生物多样性显著相关，土壤有机碳和镉离子均与AOB和nirS微生物多样性显著相关。综上所述，饲草与杂粮保护播种会影响饲草根际土壤氨氧化和反硝化微生物多样性、组成和共现网络，揭示了保护播种技术下氮素高效利用的微生物机制，且以2∶3的比例种植作物可能会发挥出保护播种技术最佳优势。

关键词: 保护播种, 氨氧化, 反硝化, 功能基因, 多样性, 共现网络

Abstract:

Companion planting of broomcorn millet （Panicum miliaceum） with alfalfa （Medicago sativa） can improve nitrogen utilization. To explore the mechanism of this effect， we investigated the impact of different planting ratios of broomcorn millet and alfalfa on ammonia-oxidizing archaea （AOA）， ammonia-oxidizing bacteria （AOB）， denitrifying microorganisms （nirK-， nirS-， and nosZ-containing microbes）， and other soil factors in alfalfa rhizosphere soil under companion planting in the dry farming area of the Loess Plateau. Four broomcorn millet-alfalfa planting ratios ［1∶1 （1P1M）， 1∶2 （1P2M）， 1∶3 （1P3M）， and 2∶3 （2P3M）］ were established with alfalfa monoculture （M） as the control. The diversity， structure， composition， and co-occurrence network of ammonia-oxidizing and denitrifying microorganisms in alfalfa rhizosphere soil were determined by high-throughput sequencing， and their correlations with soil factors were analyzed. The results showed that， as the ratio of broomcorn millet increased， the soil total nitrogen content and stable carbon and nitrogen isotope values increased， and significantly increased in 2P3M （P<0.05）； and the soil organic carbon content first decreased and then increased， with a significant decrease in 2P3M （P<0.05）. Companion planting increased the richness of ammonia-oxidizing and nosZ-containing microorganisms， but decreased the richness of nirK-containing microorganisms. The alpha diversity of AOA was sensitive to the planting ratio. However， a beta diversity analysis revealed no significant differences in ammonia-oxidizing and denitrifying community structures among the different planting ratios. Nitrososphaera and Nitrosospira were the dominant genera of AOA and AOB， respectively， accounting for over 85% of their communities based on abundance. The denitrifying microbial community was enriched in genera that promote plant growth and nutrient transformation， such as Achromobacter among the nirK-containing microorganisms and Azoarcus among the nirS-containingmicroorganisms. A co-occurrence network analysis revealed a more complex co-occurrence network of soil ammonia-oxidizing and denitrifying communities in 2P3M than in the other treatments， and this was mainly evident in the network complexity and modularity indicators. The contents of total nitrogen and calcium and magnesium ions were significantly correlated with AOA microbial diversity， and the contents of soil organic carbon and cadmium ions were significantly correlated with AOB and nirS microorganisms. In summary， companion planting affected the diversity， composition， and co-occurrence network of ammonia-oxidizing and denitrifying microorganisms in the rhizosphere soil of alfalfa， revealing the microbial mechanism of efficient nitrogen utilization under this technology. Based on these results， the broomcorn millet∶alfalfa planting ratio of 2∶3 appears to be the most effective for exploiting the advantages of companion planting.

Key words: companion planting, ammonia oxidation, denitrification, functional gene, diversity, co-occurrence network

李若璇, 李升郅粲, 陈奕彤, 孙雨豪, 杨培志, 崔彦农, 龙明秀, 何树斌. 保护播种下紫花苜蓿根际土壤氨氧化和反硝化微生物群落对糜子种植比例变化的响应[J]. 草业学报, 2025, 34(6): 110-121.

Ruo-xuan LI, Sheng-zhi-can LI, Yi-tong CHEN, Yu-hao SUN, Pei-zhi YANG, Yan-nong CUI, Ming-xiu LONG, Shu-bin HE. Effects of different planting ratios of broomcorn millet （Panicum miliaceum） on ammonia-oxidizing and denitrifying microorganisms in rhizosphere soil of alfalfa （Medicago sativa）[J]. Acta Prataculturae Sinica, 2025, 34(6): 110-121.

图/表 7

表1 Illumina测序分析中使用的扩增引物和扩增片段大小

Table 1 Amplification primers and amplification sizes used in Illumina sequencing analysis

功能基因 Functional genes	引物Primers	引物序列Primer sequences （5′—3′）	长度Length （bp）	参考文献Reference
AOA	Arch-amoA26F Arch-amoA417R	GACTACATMTTCTAYACWGAYTGGGC GGKGTCATRTATGGWGGYAAYGTTGG	417	［18］
AOB	amoA1F amoA2R	GGGGTTTCTACTGGTGGT CCCCTCKGSAAAGCCTTCTTC	500	［19］
nirK	NIRK-F NIRK-R	TCATGGTGCTGCCGCGYGANGG GAACTTGCCGGTKGCCCAGAC	400	［20］
nirS	CD3AF R3CDR	GTSAACGTSAAGGARACSGG GASTTCGGRTGSGTCTTGA	400	［21］
nosZ	NOSZ-1126F NOSZ-1381R	GGGCTBGGGCCRTTGCA GAAGCGRTCCTTSGARAACTTG	400	［22］

表2 不同保护播种比例下土壤因子变化

Table 2 Changes of soil factors in different companion planting ratios

处理 Treatments	土壤有机碳 SOC （g·kg^-1）	总氮 TN （g·kg^-1）	铵态氮 NH₄⁺-N （mg·kg^-1）	硝态氮 NO₃^--N （mg·kg^-1）	稳定碳同位素 δ¹³C（‰）	稳定氮同位素 δ¹⁵N（‰）	钙离子 Ca²⁺ （mg·kg^-1）	镁离子 Mg²⁺ （mg·kg^-1）	镉离子 Cd²⁺ （mg·kg^-1）
M	5.49±0.55a	0.27±0.07bc	4.98±0.92a	2.96±0.50a	-7.20±0.08b	26.35±2.12b	4.59±0.27c	35.83±0.01ab	0.040±0.00a
1P3M	4.18±0.87b	0.22±0.02c	4.22±0.49a	3.02±0.70a	-7.03±0.38ab	26.71±4.51ab	5.42±0.26a	39.92±2.43a	0.039±0.00ab
1P2M	4.44±0.97ab	0.25±0.05bc	4.63±1.04a	2.89±0.44a	-6.81±0.44ab	26.81±1.77ab	5.20±0.43ab	38.93±3.02ab	0.033±0.01b
2P3M	4.23±0.57b	0.41±0.04a	4.67±1.03a	2.99±0.98a	-6.62±0.22a	32.12±3.89a	4.73±0.47bc	36.27±3.43ab	0.039±0.00ab
1P1M	5.16±0.66ab	0.31±0.08b	4.52±0.83a	3.01±0.38a	-6.99±0.34ab	27.33±2.25ab	4.39±0.07c	34.84±2.57b	0.036±0.01ab

图1 不同保护播种比例下氨氧化和反硝化微生物群落α多样性AOA：氨氧化古菌 Ammonia-oxidizing archaea；AOB：氨氧化细菌 Ammonia-oxidizing bacteria；nirK：nirK微生物群落nirK-containing microorganisms；nirS：nirS微生物群落nirS-containing microorganisms；nosZ：nosZ微生物群落nosZ-containing microorganisms. 下同The same below. *， P<0.05； **， P<0.01 and ***， P<0.001.

Fig. 1 Alpha diversity of the alfalfa rhizosphere soil ammonia-oxidizing and denitrifying microbial communities in different companion planting ratios

图2 不同保护播种比例下紫花苜蓿根际土壤氨氧化和反硝化微生物群落的主坐标分析

Fig. 2 Principal coordinates analysis of the alfalfa rhizosphere soil ammonia-oxidizing and denitrifying microbial communities in different companion planting ratios

图 3 不同保护播种比例下紫花苜蓿根际土壤氨氧化和反硝化微生物群落在属水平上的群落组成

Fig. 3 Community composition at the genus levels of the alfalfa rhizosphere soil ammonia-oxidizing and denitrifying microbial communities in different companion planting ratios

图4 不同保护播种比例下氨氧化和反硝化微生物群落在共现网络的可视化网络（A）、网络组成（B）及拓扑学性质（C）节点间的连接代表它们之间有强且显著的相关性。网络（A）中的点代表OTUs。A connection represents a strong （Spearman’s ρ>0.60） and significant （P<0.05） correlation. Different dots in （A） represent the OTUs in the network.

Fig. 4 Visual network （A）， network composition （B） and topology properties （C） of microbial co-occurrence network in ammonia-oxidizing and denitrifying communities in different companion planting ratios

图5 紫花苜蓿根际氨氧化和反硝化微生物群落与土壤因子的关系热图颜色和网格大小代表了相关性大小。绿色和粉色方框分别代表正相关和负相关。线条粗细映射相关性值。节点与土壤因子之间连线的颜色具有统计学意义。Heat map color and grid size represent the size of correlation value. Green boxes represent positive correlations， and pink boxes represent negative correlations. The thickness of each line is mapped in this interval according to the correlation value. The color of the connection between the nodes and the soil factors indicate statistical significance. *， P<0.05； **， P<0.01 and ***， P<0.001.

Fig. 5 Relationship of ammonia-oxidizing and denitrifying microbial community diversity with alfalfa rhizosphere soil factors

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