Effect of elevated atmospheric CO2 concentration on the metabolic function of microbe in rhizosphere of different soybean cultivars
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摘要: 作为全球气候变化主要因子, 大气CO2浓度升高以植物为媒介间接影响土壤微生物代谢功能, 进而影响由土壤微生物参与的土壤养分循环过程和土壤碳库的平衡。本研究以不同年代培育的大豆品种: ‘小黄金’(XH)、‘牡丰5号’(MF)、‘绥农14号’ (SN)和‘东生1号’ (DS)为试验材料, 采用开顶式气室(OTC)模拟21世纪中叶大气CO2浓度(550 mol∙L−1)升高条件(EC), 并以正常大气CO2浓度为对照(CK), 通过BIOLOG方法, 解析了大气CO2浓度升高条件下大豆根际土壤微生物对不同碳源利用的特征。结果表明, 不同大豆品种根际土壤微生物群落碳源代谢特征不同, 单孔平均颜色变化率(AWCD)总体趋势表现为MF>SN>DS>XH。微生物功能多样性指数和主成分分析结果显示, 大气CO2浓度升高对不同大豆品种根际微生物碳源代谢特征的影响不一致, 其中XH、MF和DS的根际微生物功能受大气CO2浓度升高影响不显著, 而SN的根际微生物功能受大气CO2浓度升高影响显著; 主成分贡献率结果表明, SN的EC碳源变化与主成分1中正相关的碳源种类相关性较强, 而CK的碳源变化与主成分1中负相关的碳源种类相关性较强, 其中L-精氨酸和2-羟基苯甲酸为不利于植物生长的碳源类型, 未来大气CO2浓度升高是否会增加有害/有益根系分泌物的释放有待通过田间原位试验进一步佐证。综上, 未来大气CO2浓度升高对大豆根际微生物碳源代谢功能的影响与大豆品种有关, 同时, 大豆品种和大气CO2升高及其交互作用显著影响部分碳源的代谢。Abstract: Climate change, characterized by increased concentrations of CO2, can substantially stimulate plant photosynthesis and growth, and subsequently change the quantity and quality of below-ground rhizodeposition. From the perspective of soil microbiology, these alterations may indirectly induce changes in soil microbial biomass and function, and further influence the processes of soil organism evolution, such as nutrient transformation and the balance of soil carbon storage. In this study, four soybean cultivars, ‘Xiaohuangjin’ (XH), ‘Mufeng 5’ (MF), ‘Suinong 14’ (SN), and ‘Dongsheng 1’ (DS), which are released in 1951, 1972, 1996 and 2003, respectively, widely planted in Northeast China then, were selected. Soil samples were collected from the rhizosphere of soybeans grown in either ambient CO2 (410 mol∙L−1, CK) or elevated CO2 (550 mol∙L−1, EC) at the beginning of the seed stage. Each CO2 treatment was performed in triplicate. Thus, there were 12 OTCs (open top chamber) in total. The effects of elevated CO2 on microbial metabolic function were studied using the BIOLOG technique. The results showed that different soybean cultivars had different metabolic patterns and the order of average well color development (AWCD) ranked as: MF > SN > DS > XH. Biodiversity indices and PCA analysis revealed that the metabolic patterns of cultivars XH, MF, and DS were resistant to elevated CO2, while elevated CO2 significantly changed the metabolic patterns of SN. The contribution of each carbon source to the PCA indicated strong correlations between the variations of carbon sources in the EC treatment and CK of SN with the positive and negative carbon sources in PC1, respectively. In addition, some specific carbon sources in CK of SN, such as L-arginine and 2-hydroxy benzoic acid, are harmful for soybean growth, and whether elevated CO2 may increase soybean diseases resistance merits further analysis. Meanwhile, there were interaction effects between soybean cultivars and elevated CO2 on specific carbon sources. In summary, this study illustrates that different soybean cultivars have different metabolic functions and respond differently to increased CO2.
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图 1 正常CO2浓度(a)和CO2浓度升高(b)条件下不同大豆品种根际微生物在BIOLOG-ECO板上的平均颜色变化率(AWCD)
图例中各缩写代表不同大豆品种: DS, 东生1号; MF, 牡丰5号; SN, 绥农14号; XH, 小黄金。DS, MF, SN and XH stand for soybean cultivars of ‘Dongsheng 1’ ‘Mufeng 5’ ‘Suinong 14’ and ‘Xiaohuangjin’, respectively.
Figure 1. AWCD of microbe in rhizosphere of different soybean cultivars under ambient CO2 (a) and elevated CO2 (b) concentrations
图 2 不同大豆品种根际土壤微生物在大气CO2浓度升高(EC)及不升高(CK)处理下的微生物多样性指数
“ns”表示两处理间无显著差异, “**”表示两处理间差异显著。各缩写代表不同大豆品种: DS, 东生1号; MF, 牡丰5号; SN, 绥农14号; XH, 小黄金。“ns” means no significant difference between two treatments, while “**” means significant differences between two treatments. DS, MF, SN and XH stand for soybean cutivars of ‘Dongsheng 1’ ‘Mufeng 5’ ‘Suinong 14’ and ‘Xiaohuangjin’, respectively.
Figure 2. Diversity indexes of microbe in rhizosphere of different soybean cultivars under elevated CO2 (EC) and ambient CO2 (CK) concentrations
图 3 不同大豆品种根际微生物功能多样性在大气CO2浓度升高(EC)及不升高(CK)处理下的主成分分析
各缩写代表不同大豆品种: DS, 东生1号; MF, 牡丰5号; SN, 绥农14号; XH, 小黄金。H2等表示碳源。DS, MF, SN and XH stand for soybean cultivars of ‘Dongsheng 1’ ‘Mufeng 5’ ‘Suinong 14’ and ‘Xiaohuangjin’, respectively. H2, etc. stand for carbon sources.
Figure 3. Principal component analysis of microbe in rhizosphere of different soybean cultivars under elevated CO2 EC) and ambient CO2 (CK) concentrations
表 1 31种碳源在大气CO2浓度升高(EC)及不升高(CK)处理下在第1、2和3主成分上的载荷值
Table 1. Carbon substrates loaded on the principal components in analysis of BIOLOG ECO micro-plate data under elevated (EC) and ambient CO2 (CK) conditions
碳源化学类别 Carbon substrate 碳源 Carbon substrate PC1 PC2 PC3 胺类 Amines/amides 腐胺 Putrescine −0.491 −0.448 0.287 苯乙胺 Phenylethylamine −0.512 0.153 0.108 氨基酸类 Amino acids 甘氨酰-L-谷氨酸 Glycyl-L Glutamic Acid −0.544 0.038 −0.140 L-精氨酸 L-Arginine −0.551 −0.466 0.129 L-天门冬酰胺 L-Asparagine −0.595 −0.109 0.241 L-苯丙氨酸 L-Phenylalanine −0.519 0.355 −0.026 L-丝氨酸 L-Serine −0.713 −0.376 0.033 L-苏氨酸 L-Threonine −0.587 0.072 −0.467 糖类 Carbohydrates α-D-乳糖 α-D-Lactose 0.170 0.595 0.224 β-甲基-D-葡萄糖苷 β-Methyl-D Glucoside −0.732 0.343 −0.117 D-纤维二糖 D-Cellobiose −0.697 0.004 −0.332 D-甘露醇 D-Mannitol −0.835 0.173 −0.05 D-木糖/戊醛糖 D-Xylose −0.110 0.683 0.078 i-赤藓糖醇 i-Erythritol −0.606 0.107 0.208 N-乙酰-D葡萄糖氨 N-Acetyl-D Glucosamine −0.696 0.413 −0.270 羧酸类 Carboxylic acids γ-羟丁酸 γ-Hydroxybutyric Acid −0.519 −0.053 0.170 α-丁酮酸 α-Ketobutyric Acid 0.251 −0.010 0.002 4-羟基苯甲酸 4-Hydroxy Benzoic Acid −0.390 0.416 0.669 2-羟基苯甲酸 2-Hydroxy Benzoic Acid −0.066 −0.109 −0.186 D-半乳糖酸γ-内酯 D-Galactonic Acid γ-Lactone −0.537 −0.413 −0.420 D-半乳糖醛酸 D-Galacturonic Acid −0.756 −0.005 −0.226 D-葡糖胺酸 D-Glucosaminic Acid −0.570 −0.013 0.226 D-苹果酸 D-Malic Acid −0.198 −0.242 0.177 衣康酸 Itaconic Acid −0.146 −0.174 0.542 聚合物 Polymers α-环式糊精 α-Cyclodextrin −0.289 0.550 0.092 肝糖 Glycogen −0.409 0.264 −0.209 吐温40 Tween 40 −0.500 −0.368 0.312 吐温80 Tween 80 −0.741 0.112 0.267 其他类 Miscellaneous D,L-α-磷酸甘油 D,L-α-Glycerol Phosphate −0.764 −0.374 −0.156 1-磷酸葡萄糖 Glucose-1-Phosphate −0.682 0.285 −0.132 丙酮酸甲酯 Pyruvic Acid Methyl Ester −0.597 −0.231 0.183 
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表 2 大气CO2浓度、品种及两者的交互作用对大豆根际土壤微生物不同碳源利用影响的显著性分析
Table 2. Significance analysis of effects of atmospheric CO2 concentration, soybean cultivars and their interaction on the utilization of different carbon sources of microbe in rhizosphere of soybean
胺类 Amines/amides 氨基酸类 Amino acids 糖类 Carbohydrates H4 G4 F4 A4 B4 C4 D4 E4 H1 A2 G1 D2 B2 C2 E2 CO2 (C) 0.923 0.909 0.801 0.157 0.893 0.264 0.032* 0.744 0.538 0.031* 0.235 0.864 0.318 0.331 0.53 品种
Cultivar (V)0.082 0.058 0.043* 0** 0.068 0.916 0.010* 0.172 0.790 0.129 0.110 0.104 0.646 0.050 0.062 C×V 0.023* 0.134 0.146 0.026* 0.010* 0.748 0.012* 0.283 0.365 0.538 0.130 0.602 0.369 0.492 0.540 羧酸类 Carboxylic acids 聚合物 Polymers 其他类 Miscellaneous E3 G3 D3 C3 A3 B3 F2 H3 F3 E1 F1 C1 D1 H2 G2 B1 CO2 (C) 0.062 0.197 0.208 0.117 0.006** 0.522 0.677 0.783 0.047* 0.506 0.579 0.424 0.678 0.362 0.403 0.406 品种
Cultivar (V)0.628 0.070 0.177 0.688 0.121 0.034* 0.014* 0.702 0.143 0.277 0.304 0.080 0** 0.120 0.011* 0.029* C×V 0.020* 0.176 0.388 0.688 0.527 0.235 0.013* 0.075 0.022* 0.689 0.907 0.042* 0.086 0.082 0.539 0.462 **和*分别表示在P<0.01和P<0.05水平影响显著。H4: 腐胺; G4: 苯乙胺; F4: 甘氨酰-L-谷氨酸; A4: L-精氨酸; B4: L-天门冬酰胺; C4: L-苯丙氨酸; D4: L-丝氨酸; E4: L-苏氨酸; H1: α-D-乳糖; A2: β-甲基-D-葡萄糖苷; G1: D-纤维二糖; D2: D-甘露醇; B2: D-木糖/戊醛糖; C2: i-赤藓糖醇; E2: N-乙酰-D葡萄糖氨; E3: γ-羟丁酸; G3: α-丁酮酸; D3: 4-羟基苯甲酸; C3: 2-羟基苯甲酸; A3: D-半乳糖酸γ-内酯; B3: D-半乳糖醛酸; F2: D-葡糖胺酸; H3: D-苹果酸; F3: 衣康酸; E1: α-环式糊精; F1: 肝糖; C1: 吐温40; D1: 吐温80; H2: D,L-α-磷酸甘油; G2: 1-磷酸葡萄糖; B1: 丙酮酸甲酯。** and * indicate significant effects at P<0.01 and P<0.05 levels, respectively. H4: Putrescine; G4: Phenylethylamine; F4: Glycyl-L Glutamic Acid; A4: L-Arginine; B4: L-Asparagine; C4: L-Phenylalanine; D4: L-Serine; E4: L-Threonine; H1: α-D-Lactose; A2: β-Methyl-D Glucoside; G1: D-Cellobiose; D2: D-Mannitol; B2: D-Xylose; C2: i-Erythritol; E2: N-Acetyl-D Glucosamine; E3: γ-Gydroxybutyric Acid; G3: α-Ketobutyric Acid; D3: 4-Hydroxy Benzoic Acid; C3: 2-Hydroxy Benzoic Acid; A3: D-Galactonic Acid γ-Lactone; B3: D-Galacturonic Acid; F2: D-Glucosaminic Acid; H3: D-Malic Acid; F3: Itaconic Acid; E1: α-Cyclodextrin; F1: Glycogen; C1: Tween 40; D1: Tween 80; H2: D, L-α-Glycerol Phosphate; G2: Glucose-1-Phosphate; B1: Pyruvic Acid Methyl Ester.
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