Ecological stoichiometry characteristics of soil extracellular enzymes under different citrus ages and analysis of their driving factors
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摘要:
土壤胞外酶生态化学计量特征可用于评估微生物对资源和养分的获取情况, 是评价土壤肥力和微生物活性的重要指标。然而, 高强度集约化柑橘种植对土壤胞外酶计量特征的影响机制尚不清楚。为探讨集约化农业种植对土壤酶化学计量特征及其对种植年限的响应, 本研究以三峡库区河岸带不同种植年限柑橘为研究对象, 采集根际土壤, 测定与碳氮磷循环相关的胞外酶活性和功能微生物基因丰度, 通过胞外酶生态化学计量特征去评估土壤微生物对氮磷养分及资源的需求状况。结果表明, 柑橘种植年限的增加会提高土壤有效氮和有效磷的含量, 其中磷素累积更为明显。种植30 a的柑橘土壤有效磷含量约为5 a柑橘土壤的3.5倍, 远高于柑橘生长需求阈值。柑橘种植年限增加会显著降低土壤碳磷循环相关酶活性, 增加氮获取酶活性。从功能基因角度来看, 编码碱性磷酸酶的phoD基因丰度显著下降, 从4.84×107 copies·g−1下降至9.24×106 copies·g−1。土壤功能微生物基因丰度的下降是导致碱性磷酸酶活性降低的直接因素。土壤酶化学计量特征也随着柑橘种植年限的增加而改变。土壤酶矢量角度由58.21°降低至18.70°, 表明土壤微生物对养分的需求由磷限制转换为氮限制, 低柑橘种植年限土壤微生物以磷限制为主, 高柑橘年限土壤微生物以氮限制为主。高强度柑橘种植过程中, 需减少磷肥施用, 增加有机肥等碳源投入, 提高微生物活性。研究结果可为高强度集约化柑橘种植土壤质量提升和果园的可持续管理提供理论依据。
Abstract:The ecological stoichiometric characteristics of soil extracellular enzymes can be used to evaluate the acquisition of resources and nutrients by microorganisms, sensitively reflect the metabolic characteristics of soil microorganisms, and are important indicators to evaluate soil fertility and microbial activity. Intensive agriculture is characterized by the long-term application of large amounts of chemical fertilizers, which often leads to changes in the composition and activity of microbial communities in the soil. However, the mechanism by which high-intensity citrus cultivation affects the stoichiometric characteristics of soil extracellular enzymes remains unclear. To investigate the effects of intensive agricultural cultivation on the stoichiometric characteristics of soil enzymes and their responses to planting years, this study focused on citrus orchards with varying planting years in the riparian zones of the Three Gorges Reservoir. Rhizosphere soils were collected to measure extracellular enzyme activity related to carbon, nitrogen, and phosphorus cycling. The ecological stoichiometry of extracellular enzymes was used to assess the demand for nitrogen, phosphorus, and other resources from soil microorganisms. The results indicated that with increasing citrus planting years, the content of available nitrogen and phosphorus in the soil increased, particularly with significant accumulation of soil phosphorus. The soil phosphorus content in 30-year citrus orchards was 2.5 times higher than that in 5-year orchards, far exceeding the threshold required for citrus growth. Additionally, in citrus soils with long citrus planting years, other forms of phosphorus also accumulate in large amounts. In citrus soils with different planting years, enzyme activities related to the acquisition of carbon, nitrogen, and phosphorus was different. Increasing the citrus planting year significantly reduced the activity of soil enzymes related to carbon and phosphorus cycling, while increasing nitrogen acquisition enzyme activity. From a functional gene perspective, the abundance of the phoD gene encoding alkaline phosphatase significantly decreased from 4.84×107 copies·g−1 to 9.24×106 copies·g−1. Decrease in the abundance of soil functional microbial genes directly contribute to the reduction in the alkaline phosphatase activity. The stoichiometric characteristics of soil enzymes also changed with increasing citrus planting duration, and the enzyme vector model revealed that the soil enzyme vector angle decreased from 58.21° to 18.70°, indicating a transition in soil microbial nutrient demand from phosphorus to nitrogen limitation. Soil microbial communities in the 5-year citrus orchards were primarily P-limited, whereas those in the 30-year orchards were primarily N-limited. In the process of high-intensity citrus planting, it is necessary to reduce the application of phosphate fertilizers and increase the input of carbon sources to promote phosphorus utilization through carbon and alleviate microbial nutrient limitations. Alkaline conditioners should be appropriately added to soils with high citrus planting years to alleviate soil acidification. These findings provide a theoretical basis for improving soil quality and sustainable management of orchards under intensive citrus cultivation.
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图 1 不同柑橘种植年限(5 a和30 a)土壤胞外酶之间的回归关系
BG: β-1, 4-葡萄糖苷酶; CBH: 纤维二糖水解酶; NAG: β-N-乙酰氨基葡萄糖苷酶; ALP: 碱性磷酸酶。BG: β-1,4-glucosidase; CBH: cellobiohydrolase; NAG: β-N-acetylglucosaminidase; ALP: alkaline phosphatase.
Figure 1. Regression relationship between extracellular enzymes in soil under different planting years (5 a and 30 a) of citrus
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图 2 不同种植年限(5 a和30 a)土壤胞外酶矢量特征
C: 碳获取酶活性; N: 氮获取酶活性; P: 磷获取酶活性。C: carbon acquisition enzyme activity; N: nitrogen acquisition enzyme activity; P: phosphorus acquisition enzyme activity.
Figure 2. Vector characteristics of soil extracellular enzymes under different planting years (5 a and 30 a) of citrus
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图 3 土壤胞外酶与土壤基本特性、磷组分及phoD基因丰度的相关性
红色圆圈表示正相关, 蓝色圆圈表示负相关; 圆圈大小表示相关性大小。AVP: 有效磷; BG: β-1,4-葡萄糖苷酶; CBH: 纤维二糖水解酶; NAG: β-N-乙酰氨基葡萄糖苷酶; ALP: 碱性磷酸酶; CaCl2-P: 可交换态磷; Citrate-P: 根系分泌有机酸溶解态磷; HCl-P: 矿物结合态磷。Red and blue circles refer to positive and negative correlation, respectively. Circle size refers to the correlation. AVP: available phosphorus; BG: β-1,4-glucosidase; CBH: cellobiohydrolase; NAG: β-N-acetylglucosaminidase; ALP: alkaline phosphatase; CaCl2-P: exchangeable phosphorus; Citrate-P: organic acids dissolved phosphorus of root secretion; HCl-P: mineral bound phosphorus.
Figure 3. Correlation between extracellular enzymes in soil and basic soil characteristics, phosphorus components, and abundance of phoD genes
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图 4 酶矢量角度与有效磷、有效氮和phoD基因丰度之间的回归关系
AVP: 有效磷。AVP: available phosphorus.
Figure 4. Regression relationship between enzyme vector angle and available phosphorus, available nitrogen and phoD gene abundance
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图 5 长时间柑橘种植改变土壤微生物对养分的获取策略
AVP: 有效磷; ALP: 碱性磷酸酶。AVP: available phosphorus; ALP: alkaline phosphatase.
Figure 5. Long-term citrus cultivation changes nutrient acquisition strategies of soil microorganisms
表 1 土壤胞外酶及其编号与底物
Table 1 Basic information of soil extracellular enzymes
酶 Enzyme 缩写 Abbreviation 编号 Code 底物 Substrate β-1,4-葡萄糖苷酶
β-1,4-glucosidaseBG 3.2.1.21 4-MUB-β-D-葡萄糖苷
4-MUB-β-D-glucoside纤维二糖水解酶
CellobiohydrolaseCBH 3.2.1.91 4-MUB-β-D-纤维二糖苷
4-MUB-β-D-cellobiosideβ-N-乙酰氨基葡萄糖苷酶
β-N-acetylglucosaminidaseNAG 3.1.6.1 4-MUB-N-乙酰基-b-D-氨基葡萄糖
4-MUB-N-acetyl-b-D-glucosaminide碱性磷酸酶
Alkaline phosphataseALP 3.1.3.1 4-MUB-磷酸盐
4-MUB-phosphate
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表 2 不同种植年限柑橘土壤基本理化性质
Table 2 Basic physical and chemical properties of citrus soil with different planting years
种植年限
Planting year (a)pH SOM
(g·kg−1)NO3−-N
(mg·kg−1)NH4+-N
(mg·kg−1)AVP
(mg·kg−1)CaCl2-P
(mg·kg−1)Citrate-P
(mg·kg−1)HCl-P
(mg·kg−1)5 5.75±0.75a 17.95±6.65a 37.37±36.09b 4.56±2.46b 31.40±21.16b 5.00±3.83b 216.97±242.84a 554.32±406.88b 30 3.69±0.37b 20.86±4.73a 117.34±60.53a 58.90±41.90a 108.55±26.51a 28.44±13.95a 331.06±89.93a 801.90±212.36a 同列不同小写字母表示同一指标不同柑橘年限间差异显著(P<0.05)。AVP: 有效磷; CaCl2-P: 可交换态磷; Citrate-P: 根系分泌有机酸溶解态磷; HCl-P: 矿物结合态磷。Different lowercase letters in the same column indicate significant differences between different planting years of the same indicator (P<0.05). AVP: available phosphorus; CaCl2-P: exchangeable phosphorus; Citrate-P: organic acids dissolved phosphorus of root secretion; HCl-P: mineral bound phosphorus.
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表 3 不同种植年限柑橘土壤酶活性及酶矢量
Table 3 Soil enzyme activities and enzyme vectors of citrus with different planting years
种植年限
Planting year (a)BG
(nmol·g−1·h−1)CBH
(nmol·g−1·h−1)NAG
(nmol·g−1·h−1)ALP
(nmol·g−1·h−1)角度
Angle (°)长度
Length5 67.99±44.91a 16.60±9.93a 21.26±18.60b 85.09±47.10a 58.21±10.92a 0.96±0.14a 30 22.53±14.47b 4.95±4.31b 85.16±80.77a 2.13±2.75b 18.70±10.20b 1.00±0.08a 同列不同小写字母表示同一指标不同柑橘年限间差异显著(P<0.05)。BG: β-1,4-葡萄糖苷酶; CBH: 纤维二糖水解酶; NAG: β-N-乙酰氨基葡萄糖苷酶; ALP: 碱性磷酸酶。Different lowercase letters in the same column indicate significant differences between different planting years of the same indicator (P<0.05). BG: β-1,4-glucosidase; CBH: cellobiohydrolase; NAG: β-N-acetylglucosaminidase; ALP: alkaline phosphatase.
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