Effects of Straw-returning on N2O Emission and Nitrifying/Denitrifying Microbes in Paddy Soil in Southern China
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摘要:
目的 以我国南方典型赤红壤水稻土为研究对象，探究不同秸秆还田量对双季稻区晚稻季土壤N2O排放特征及硝化和反硝化微生物的影响，旨在为南方稻田N2O减排提供科学依据。 方法 以2015年设置的定位试验为研究平台，设计5个处理：（1）CK，化肥+无秸秆还田；（2）CKS，化肥+当季秸秆全量还田；（3）S10，化肥＋当季秸秆全量还田+秸秆替代10%钾肥；（4）S20，化肥＋当季秸秆全量还田+秸秆替代20%钾肥；（5）S30，化肥＋当季秸秆全量还田+秸秆替代30%钾肥。采用密闭静态暗箱-气相色谱法及宏基因组测序对气体和土壤微生物进行检测。 结果 稻田土壤N2O排放主要集中在水稻分蘖期；较CK处理，秸秆还田各处理显著降低了稻田土壤N2O累计排放量，其中，S30处理的N2O累计排放量最低，为0.09 kg·hm−2，其全球增温潜势也最低；硝化过程中，氨氧化细菌（amoA和amoB）在分蘖期和成熟期的优势菌属均为甲基孢囊菌属；反硝化过程中，nirK型反硝化细菌的芽单胞菌属、罗河杆菌属、丰祐菌属在分蘖期和成熟期细菌属中均占据主导地位；nirS型反硝化细菌的嗜甲基菌属和甲基营养型反硝化菌属在分蘖期和成熟期细菌属中均占据主导地位；分蘖期，土壤N2O排放量与nirS型反硝化细菌的甲基营养型反硝化菌属呈显著负相关、与nirS型反硝化细菌的沙壤土杆菌属呈显著正相关；成熟期，土壤N2O排放量与nirS型反硝化细菌的水生细菌属呈极显著正相关关系。 结论 秸秆还田显著降低了稻田土壤N2O排放，AOB（amoA）和AOB（amoB）的甲基孢囊菌属是氨氧化过程的优势菌属，nirK型反硝化细菌的芽单胞菌属、罗河杆菌属、丰祐菌属，和nirS型反硝化细菌的嗜甲基菌属及甲基营养型反硝化菌属是反硝化过程的优势菌属。 Abstract:Objective Effects of returning spent straws to soil after crop harvest as a conditioner/fertilizer on the N2O emission and nitrifying/denitrifying microbes on a double-cropping rice field in late season were studied. Method Based on a positioning experiment set up in 2015, the field soil was treated with (1) chemical fertilizer without straw-returning (CK), (2) chemical fertilizer + 100% straw-returning in same season (CKS), (3) CKS + straws to replace 10% potassium fertilizer (S10), (4) CKS + straws to replace 20% potassium fertilizer (S20) or (5) CKS + straws to replace 30% potassium fertilizer (S30). N2O emitted and microorganisms in the soils were detected using the closed static dark box-gas chromatography and metagenomic sequencing technique. Result N2O was released from the paddy soils basically during the rice tillering stage. Compared with CK, returning spent straws to the ground significantly reduced the cumulative gas emission from the soil under the S30 treatment, which rendered the lowest rate at 0.09 kg·hm-2, and thus, the least contribution to global warming. Of the nitrifying microflora in soil, the dominant genus of ammonia-oxidizing bacteria (i.e., amoA and amoB) at the tillering and mature stages was Methylospora. Among the denitrifying microbes, the nirK-type Gemmatimonadetes, Rhodobacter, and Opitutus and the nirS-type Methylobacillus and Methylotenera were dominantly present. At the rice tillering stage, the N2O emissions were significantly inversely correlated with Methylotenera but positively with Ramlibacter. At the mature stage, a significant correlation between the soil N2O emission and Aquabacterium population was observed. Conclusion By returning the spent straws to the field, a significant reduction on the N2O emissions from paddy soil was resulted. And Methylospora was the dominant genus involved in the ammonia oxidation, whereas the nirK-type Gemmatimonadetes,Rhodobacter and Opitutus and the nirS-type Methylobacillus and Methylotenera were the dominant genera of denitrifying bacteria. -
图 1 水稻四个生长阶段稻田土壤N2O排放通量
Figure 1. N2O emission flux of paddy soil at 4 rice growth stages
图 2 稻田土壤分蘖期和成熟期N2O排放通量
不同小写字母表示不同处理间差异显著（P＜0.05），图3、表3同。
Figure 2. N2O emission flux of paddy soil at rice tillering and mature stages
The different letters indicates significant difference among treatments at P＜0.05 level. Same for Figure 3 and Table 3.
图 3 水稻生育期稻田土壤N2O累计排放量
Figure 3. Cumulative N2O emission from paddy soil in rice growth period
图 4 秸秆还田下土壤硝化和反硝化微生物群落非度量多维尺度分析（NMDS）
Figure 4. Non-metric multidimensional scale analysis on soil nitrifying/denitrifying microbes under straw-returning practice
图 5 秸秆还田下硝化细菌属相对丰度
Figure 5. Relative abundance of nitrifying microbes under straw-returning practice
图 6 秸秆还田下反硝化细菌属相对丰度
Figure 6. Relative abundance of denitrifying microbes under straw-returning practice
表 1 不同处理化肥养分投入量
Table 1. Nutrients in treatment fertilizers
（kg·hm−2） 处理
Treatment早稻 Early rice 晚稻 Late rice 总计 Total N P2O5 K2O N P2O5 K2O N P2O5 K2O CK 155.3 47.3 135 155.3 47.3 135 310.5 94.5 270 CKS 155.3 47.3 135 155.3 47.3 135 310.5 94.5 270 S10 155.3 47.3 121.5 155.3 47.3 121.5 310.5 94.5 243 S20 155.3 47.3 108 155.3 47.3 108 310.5 94.5 216 S30 155.3 47.3 94.5 155.3 47.3 94.5 310.5 94.5 189 
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表 2 硝化反硝化过程关键功能基因
Table 2. Key functional genes involving nitrification and denitrification in soil
功能基因分组
Functional gene grouping目标基因
Target geneKEGG编号
KEGG number基因功能详情
Gene Function Details硝化作用 Nitrification amoA（AOB） K10944 氨单加氧酶亚基A Ammonia monooxygenase subunit A amoB（AOB） K10945 氨单加氧酶亚基B Ammonia monooxygenase subunit B 反硝化作用 Denitrification nirK K00368 亚硝酸还原酶 Nitrite reductase nirS K15864 亚硝酸还原酶 Nitrite reductase
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表 3 不同处理下土壤N2O的增温潜势
Table 3. Global warming potentials due to N2O emitted from soils under treatments
（kg·hm-2） 处理 Treatement CK S10 S20 S30 CKS GWP 204.05±12.24 a 31.80±3.06 d 95.40±9.18 c 23.85±1.53 d 121.90±9.18 b
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表 4 分蘖期土壤N2O排放通量与硝化反硝化微生物群落相关性
Table 4. Correlation between soil N2O emission and nitrifying/denitrifying microbes at rice tillering stage
硝化和反硝化功能基因
Nitrification and Denitrifying gene菌属（属水平）
Bacteria(genus level)相关性系数
Correlation coefficientP值
P−valueamoA （AOB） 甲基孢囊菌属（Methylocystis） 0.107 0.703 亚硝化螺菌属（Nitrosospira） −0.068 0.809 硝化螺菌属（Nitrospira） 0.227 0.416 amoB（AOB） 慢生根瘤菌属（Bradyrhizobium） −0.038 0.892 甲基孢囊菌属（Methylocystis） −0.152 0.588 nirK 气火菌属（Aeropyrum） −0.098 0.729 Ardenticatena −0.250 0.370 慢生根瘤菌属（Bradyrhizobium） 0.078 0.781 厌氧铵氧化菌（Candidatus Brocadia） −0.236 0.396 戴氏菌属（Dyella） −0.018 0.948 芽单胞菌属（Gemmatimonas） 0.260 0.348 新草螺菌属（Noviherbaspirillum） −0.118 0.676 丰祐菌属（Opitutus） −0.489 0.064 生丝微菌属（Pseudolabrys） −0.225 0.419 罗尔斯顿菌属（Ralstonia） −0.136 0.628 罗河杆菌属（Rhodanobacter） −0.073 0.795 nirS 水生细菌（Aquabacterium） −0.136 0.629 广泛固氮氢自养单胞菌（Azohydromonas） −0.117 0.677 慢生根瘤菌属（Bradyrhizobium） −0.210 0.453 累积念珠菌（Candidatus Accumulibacter） −0.057 0.839 草螺菌（Herbaspirillum） 0.048 0.864 嗜甲基菌属（Methylobacillus） −0.166 0.555 甲基营养型反硝化菌（Methylotenera） −0.606* 0.017 新草螺菌属（Noviherbaspirillum） −0.384 0.158 沙壤土杆菌（Ramlibacter） 0.521* 0.046 红长命菌属（Rubrivivax） −0.110 0.698 硫氧化菌（Sulfurovum） 0.229 0.412 陶厄氏菌（Thauera） 0.120 0.670 硫杆菌（Thiobacillus） 0.248 0.372 辫硫细菌属（Thioploca） 0.030 0.915 “*”和“**”分别表示相关性为显著（P＜0.05）和极显著（P＜0.01）。下同。
* and ** indicate significant correlation at P＜0.05, and extremely significant at P＜0.01, respectively. Same for below.
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表 5 成熟期土壤N2O排放通量与硝化反硝化微生物群落相关性
Table 5. Correlation between soil N2O emission and nitrifying/denitrifying microbes at rice mature stage
硝化和反硝化功能基因
Nitrification and Denitrifying gene菌属（属水平）
Bacteria (genus level)相关性系数
Correlation coefficientP值
P−valueamoA（AOB） 甲基孢囊菌属（Methylocystis） −0.274 0.323 亚硝化螺菌属（Nitrosospira） 0.089 0.753 硝化螺菌属（Nitrospira） −0.098 0.729 amoB（AOB） 慢生根瘤菌属（Bradyrhizobium） −0.390 0.151 甲基孢囊菌属（Methylocystis） 0.354 0.195 nirK Ardenticatena 0.253 0.363 慢生根瘤菌属（Bradyrhizobium） 0.201 0.473 厌氧铵氧化菌（Candidatus Brocadia） −0.008 0.979 戴氏菌属（Dyella） 0.087 0.757 芽单胞菌属（Gemmatimonas） 0.005 0.987 微单孢菌属（Gemmatirosa） −0.261 0.347 新草螺菌属（Noviherbaspirillum） 0.482 0.069 丰祐菌属（Opitutus） −0.099 0.727 生丝微菌属（Pseudolabrys） −0.280 0.311 罗尔斯顿菌属（Ralstonia） 0.297 0.283 罗河杆菌属（Rhodanobacter） 0.096 0.735 nirS 水生细菌（Aquabacterium） 0.642** 0.010 广泛固氮氢自养单胞菌（Azohydromonas） −0.119 0.674 慢生根瘤菌属（Bradyrhizobium） −0.087 0.757 累积念珠菌（Candidatus Accumulibacter） −0.377 0.166 草螺菌（Herbaspirillum） −0.435 0.105 嗜甲基菌属（Methylobacillus） 0.306 0.267 甲基营养型反硝化菌（Methylotenera） 0.173 0.539 新草螺菌属（Noviherbaspirillum） −0.123 0.661 沙壤土杆菌（Ramlibacter） 0.272 0.326 硫氧化菌（Sulfurovum） 0.327 0.234 陶厄氏菌（Thauera） 0.189 0.501 辫硫细菌属（Thioploca） 0.327 0.234
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