大豆根瘤固氮特性与影响因素的研究
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摘要
大豆是我国重要的粮食和油料作物,根瘤菌通过与大豆共生结瘤固氮,为大豆生长发育提供氮素营养,研究大豆根瘤固氮特性,及肥力和水分条件对根瘤固氮的影响,为科学合理施肥,提高产量,改善品质,降低生产成本,减轻环境污染具有重要意义。
本试验于2009—2011年进行,应用15N示踪技术,采用框栽和砂培相结合的方法。框栽试验,选用不同品种大豆,利用DGGE和AFLP方法分析了与供试品种共生的根瘤菌差异,同时利用15N示踪技术研究了不同大豆品种根瘤固氮的差异；以绥农14为试验材料,利用15N示踪技术研究了肥力因素和水分条件对大豆根瘤固氮和产量的影响。砂培试验,选用绥农14为试验材料,利用15N示踪技术研究了磷素营养和钾素营养对大豆根瘤固氮和产量的影响。结果表明：
与供试大豆品种共生的根瘤菌主要是未被培养的根瘤菌。利用DGGE方法研究了龙选1号、丰收10、绥农14、小金黄和秣食豆R4期根瘤菌的差异,其差异性体现在条带数量和条带的亮度上；对条带进行测序,并与NCBI数据库相似序列进行比对,与来自环境样品的未被培养的细菌DNA序列相似度达到94%-100%。
主栽的大豆品种,黑河41、绥农14、黑农40和秣食豆AFLP结果表明,5对引物共扩增出578个位点,平均多态性位点百分率为68.86%；绥农14R2期和绥农14R6期的根瘤菌划分为一类,其余的划分为一类；不同生育时期干旱处理条件下,根瘤菌没有明显差异。
生育期长的大豆品种在根瘤固氮方面具有明显优势,供试品种根瘤固氮量和固氮率均表现为：晚熟品种>中熟品种>早熟品种,而且根瘤固氮量达到显著差异水平。
氮肥水平对大豆根瘤固氮的影响因土壤肥力而异。在速效氮含量为18.17mg-kg-1'的土壤条件下,大豆根瘤固氮量和根瘤固氮率均随氮肥水平的升高而呈单峰曲线变化,硫酸铵施用量为225kg/hm2(纯氮：47.73kg/hm2)时最有利于大豆根瘤固氮作用；在速效氮含量为50.38mg·kg-1的土壤条件下,硫酸铵施用量为75kg/hm2(纯氮：15.91kg/hm2)时最有利于大豆根瘤固氮作用,提高氮肥水平显著抑制了根瘤固氮作用。
磷素营养对大豆根瘤固氮有明显影响。砂培条件下,大豆植株根瘤固氮量和固氮率,随着磷素水平的提高均呈单峰曲线变化,当磷素浓度达到11mg/L时,再提高磷水平对根瘤固氮率促进作用不显著,当磷素浓度达到21mg/L时,再提高磷素浓度对根瘤固氮量无明显的促进作用；V3-R1、R1-R5期间断磷严重抑制大豆植株根瘤固氮量和固氮率,而R5-R7期间断磷无明显影响,表明V3-R5期间是大豆氮代谢对磷素营养的敏感期,R5以后对磷素营养不敏感；在土壤速效磷含量为26.31mg-kg-1的框栽条件下,施用磷肥有利于提高大豆根瘤固氮量,但是当重过磷酸钙施用量达到150kg/hm2(P205：69kg/hm2)时,再提高磷肥用量对大豆根瘤固氮量促进作用不显著；大豆根瘤固氮率随着磷肥施用量的增加整体呈上升趋势,当重过磷酸钙施用量达到450kg/hm2(P205：207kg/hm2)时,明显促进了全株和籽粒的根瘤固氮率。
钾素营养对大豆根瘤固氮也有一定影响。砂培条件下,大豆植株根瘤固氮量和固氮率,随着钾素营养水平的提高均呈单峰曲线变化,当钾素水平达到28mg/L时,能够满足根瘤固氮作用对钾素的需求；不同生育时期断钾对大豆植株根瘤固氮量和固氮率没有显著影响；在土壤速效钾含量为174.75mg·kg-1的框栽条件下,施钾肥促进了大豆植株根瘤固氮量和固氮率,当硫酸钾施用量达到90kg/hm2(K2O：45kg/hm2)时,能够满足根瘤固氮作用对钾素的需求。
供试大豆品种合农60、嫩丰18和绥农14的根瘤固氮量和固氮率均随生育期间水分的增加整体呈上升趋势,根瘤固氮率对水分的敏感性较根瘤固氮量低；大豆生育期间阶段性干旱对根瘤固氮的影响没有明显的规律性。
肥水因素会影响大豆产量。(1)两种供试土壤条件下,大豆产量均随施氮量的增加呈单峰曲线变化,表明氮肥水平过高和过低都会造成产量的下降。(2)砂培条件下,大豆产量随着磷素水平的提高而呈单峰曲线变化,31mg/L磷素浓度达到最大值,当磷素浓度达到21mg/L时,再提高磷素浓度无明显增产作用；V3-R1、R1-R5期间断磷显著降低了大豆产量,R5-R7期间断磷无显著影响；在十壤速效磷含量为26.31mg·kg-1的框栽条件下,施用磷肥无明显增产作用。(3)砂培条件下,大豆产量随着钾素水平的升高而呈单峰曲线变化,64mg/L钾素浓度达到最大值,当钾素浓度达到52mg/L时,再提高钾素浓度增产作用不显著；V3-R1和R1-R5断钾显著降低了大豆产量,R5-R7断钾未达到显著差异水平；在土壤速效钾含量为174.75mg·kg-1的框栽条件下,大豆产量随钾水平的升高呈上升趋势。(4)供试大豆品种合农60、嫩丰18和绥农14的产量均随着生育期间水分的增加而提高,表明提高降水量可以达到增产的目的；大豆R4-R7对干旱最敏感,此时期干旱导致产量明显下降。
Soybean is grain and oil crop in China. Nitrogen nutrition was provided for soybean growth by soybean-rhizobium symbiosis, studying on characteristics of nodule nitrogen fixation and influencing factors-fertility and moisture conditions are profound implications for appropriate application of fertilizer, increasing yield, improving quality, reduction in production costs and alleviation of environmental pollution.
This work was conducted from2009to2011, using15N tracer technique with frame tests (pot without bottom) and sand culture. In frame experiment, DGGE and AFLP methods were used to study differences of rhizobium and nodule nitrogen fixation among different soybean varieties; effect of fertility factors and moisture conditions on nodule nitrogen fixation and yield were studied with SN14by15N tracer technique. In sand culture, effect of phosphorus nutrient and potassium nutrient on nodule nitrogen fixation and yield were researched with SN14by15N tracer technique. The results showed that:
Symbiotic rhizobium with tested soybean varieties were mainly uncultured bacterium. DGGE method was used to research differences of R4rhizobium among LX1, FS10, SN14, XJH and MSD, the number and brightness of bands were different; sequence analysis of the bands with NCBI stated that the similarity was94%-100%with uncultured bacterium from environmental samples.
AFLP results of HH41, SN14, HN40and MSD showed that there were578bands with5primer combinations, the percentage of polymorphic loci was68.86%; Rhizobium could be divided into two clusters, rhizobium of SN14R2and SN14R6were one kind, others were another kind; there was no difference of rhizobium under the condition of drought at different growth stages.
Soybean varieties with the long growth period had obvious advantages on nodule nitrogen fixation, the accumulation and ratio of nodule nitrogen fixation of tested varieties both showed that late varieties>medium varieties>early varieties, and the accumulation of nodule nitrogen fixation had significant difference.
The effect of nitrogen levels on nodule nitrogen fixation of soybean was different because of soil fertility. The accumulation and ratio of nodule nitrogen fixation of soybean presented a single peak curve with nitrogen levels when the available nitrogen content of soil was18.17mg·kg-1, nodule nitrogen fixation was the highest when the amount of (NH4)2SO4was225kg/hm2(pure N:47.73kg/hm2); nodule nitrogen fixation was the highest when the amount of (NH4)2SO4was75kg/hm2(pure N:15.91kg/hm2) under the condition of available nitrogen content of soil was 50.38mg·kg-1, nodule nitrogen fixation was significantly inhibited with the nitrogen levels increasing.
There was significant effect of phosphorus nutrient on nodule nitrogen fixation of soybean. With sand culture, the accumulation and ratio of nodule nitrogen fixation of soybean presented a single peak curve with phosphorus levels, when phosphorus concentration was11mg/L, increasing phosphorus concentration again had no obvious promotion of ratio of nodule nitrogen fixation, when phosphorus concentration was21mg/L, increasing phosphorus concentration again had no obvious promotion of accumulation of nodule nitrogen fixation; accumulation and ratio of nodule nitrogen fixation decreased significantly when interruption of phosphorus supply during V3-R1and R1-R5period, while interruption of phosphorus supply during R5-R7period had no significant effect on those, which means that nitrogen metabolism of soybean was sensitive to phosphorus requirement during V3-R5period and not sensitive after R5; with frame, phosphate application could improve the accumulation of nodule nitrogen fixation of soybean when available phosphorus content of soil was26.31mg·kg-1, while the promotion effect on accumulation of nodule nitrogen fixation of soybean was not significant with phosphate fertilizer increasing when the amount of triple superphosphate was150kg/hm2(P2O5:69kg/hm2); the ratio of nodule nitrogen fixation of soybean showed an increasing tendency with phosphate fertilizer increasing, the ratio of nodule nitrogen fixation of whole plant and seeds was significantly increased when the amount of triple superphosphate was450kg/hm2(P2O5:207kg/hm2).
There was effect of potassium nutrient on nodule nitrogen fixation of soybean. With sand culture, the accumulation and ratio of nodule nitrogen fixation of soybean presented a single peak curve with potassium levels, nodule nitrogen fixation could be satisfied when potassium levels was21mg/L; there was no significant effect of interruption of potassium supply during different periods on the accumulation and ratio of nodule nitrogen fixation; potassium fertilizer could improve the accumulation and ratio of nodule nitrogen fixation of soybean when available potassium content of soil was174.75mg·kg-1, potassium could satisfy the need of nodule nitrogen fixation when the amount of K2SO4was90kg/hm2(K2O:45kg/hm2).
The accumulation and ratio of nodule nitrogen fixation showed an increasing tendency with moisture increasing in HN60, NF18and SN14, the sensitivity of the ratio of nodule nitrogen fixation to moisture was lower than the accumulation of nodule nitrogen fixation; there was no obvious rules of nodule nitrogen fixation under the condition of phased drought during growth period.
Fertility and moisture factors could influence soybean yield.(1) Soybean yield presented a single peak curve with the increase of nitrogen rate under the condition of two soil treatments, which showed that the yield decreased either too high or too low.(2) With sand culture, soybean yield presented a single peak curve with phosphorus levels, the yield reached its maximum when phosphorus concentration was31mg/L, while increasing phosphorus concentration again had no obvious promotion of yield when phosphorus concentration was21mg/L; soybean yield decreased significantly when interruption of phosphorus supply during V3-R1and R1-R5period, while interruption of phosphorus supply during R5-R7period had no significant effect on yield; soybean yield was not significantly increased with phosphate application when available phosphorus content of soil was26.31mg·kg-1.(3) Soybean yield presented a single peak curve with potassium levels increasing with sand culture, the yield reached its maximum when potassium concentration was64mg/L, while increasing potassium concentration again had no obvious promotion of yield when potassium concentration was52mg/L; soybean yield decreased significantly when interruption of potassium supply during V3-R1and R1-R5period, while interruption of potassium supply during R5-R7period had no significant effect on yield; soybean yield showed an increasing tendency with potassium levels increasing when available potassium content of soil was174.75mg·kg-1.(4) The yield of HN60, NF18and SN14was increased with moisture improving, which showed that improving precipitation was beneficial to yield; podding-seed filling period of soybean was the most sensitive to drought, and the yield was significantly decreased.
本试验于2009—2011年进行,应用15N示踪技术,采用框栽和砂培相结合的方法。框栽试验,选用不同品种大豆,利用DGGE和AFLP方法分析了与供试品种共生的根瘤菌差异,同时利用15N示踪技术研究了不同大豆品种根瘤固氮的差异；以绥农14为试验材料,利用15N示踪技术研究了肥力因素和水分条件对大豆根瘤固氮和产量的影响。砂培试验,选用绥农14为试验材料,利用15N示踪技术研究了磷素营养和钾素营养对大豆根瘤固氮和产量的影响。结果表明：
与供试大豆品种共生的根瘤菌主要是未被培养的根瘤菌。利用DGGE方法研究了龙选1号、丰收10、绥农14、小金黄和秣食豆R4期根瘤菌的差异,其差异性体现在条带数量和条带的亮度上；对条带进行测序,并与NCBI数据库相似序列进行比对,与来自环境样品的未被培养的细菌DNA序列相似度达到94%-100%。
主栽的大豆品种,黑河41、绥农14、黑农40和秣食豆AFLP结果表明,5对引物共扩增出578个位点,平均多态性位点百分率为68.86%；绥农14R2期和绥农14R6期的根瘤菌划分为一类,其余的划分为一类；不同生育时期干旱处理条件下,根瘤菌没有明显差异。
生育期长的大豆品种在根瘤固氮方面具有明显优势,供试品种根瘤固氮量和固氮率均表现为：晚熟品种>中熟品种>早熟品种,而且根瘤固氮量达到显著差异水平。
氮肥水平对大豆根瘤固氮的影响因土壤肥力而异。在速效氮含量为18.17mg-kg-1'的土壤条件下,大豆根瘤固氮量和根瘤固氮率均随氮肥水平的升高而呈单峰曲线变化,硫酸铵施用量为225kg/hm2(纯氮：47.73kg/hm2)时最有利于大豆根瘤固氮作用；在速效氮含量为50.38mg·kg-1的土壤条件下,硫酸铵施用量为75kg/hm2(纯氮：15.91kg/hm2)时最有利于大豆根瘤固氮作用,提高氮肥水平显著抑制了根瘤固氮作用。
磷素营养对大豆根瘤固氮有明显影响。砂培条件下,大豆植株根瘤固氮量和固氮率,随着磷素水平的提高均呈单峰曲线变化,当磷素浓度达到11mg/L时,再提高磷水平对根瘤固氮率促进作用不显著,当磷素浓度达到21mg/L时,再提高磷素浓度对根瘤固氮量无明显的促进作用；V3-R1、R1-R5期间断磷严重抑制大豆植株根瘤固氮量和固氮率,而R5-R7期间断磷无明显影响,表明V3-R5期间是大豆氮代谢对磷素营养的敏感期,R5以后对磷素营养不敏感；在土壤速效磷含量为26.31mg-kg-1的框栽条件下,施用磷肥有利于提高大豆根瘤固氮量,但是当重过磷酸钙施用量达到150kg/hm2(P205：69kg/hm2)时,再提高磷肥用量对大豆根瘤固氮量促进作用不显著；大豆根瘤固氮率随着磷肥施用量的增加整体呈上升趋势,当重过磷酸钙施用量达到450kg/hm2(P205：207kg/hm2)时,明显促进了全株和籽粒的根瘤固氮率。
钾素营养对大豆根瘤固氮也有一定影响。砂培条件下,大豆植株根瘤固氮量和固氮率,随着钾素营养水平的提高均呈单峰曲线变化,当钾素水平达到28mg/L时,能够满足根瘤固氮作用对钾素的需求；不同生育时期断钾对大豆植株根瘤固氮量和固氮率没有显著影响；在土壤速效钾含量为174.75mg·kg-1的框栽条件下,施钾肥促进了大豆植株根瘤固氮量和固氮率,当硫酸钾施用量达到90kg/hm2(K2O：45kg/hm2)时,能够满足根瘤固氮作用对钾素的需求。
供试大豆品种合农60、嫩丰18和绥农14的根瘤固氮量和固氮率均随生育期间水分的增加整体呈上升趋势,根瘤固氮率对水分的敏感性较根瘤固氮量低；大豆生育期间阶段性干旱对根瘤固氮的影响没有明显的规律性。
肥水因素会影响大豆产量。(1)两种供试土壤条件下,大豆产量均随施氮量的增加呈单峰曲线变化,表明氮肥水平过高和过低都会造成产量的下降。(2)砂培条件下,大豆产量随着磷素水平的提高而呈单峰曲线变化,31mg/L磷素浓度达到最大值,当磷素浓度达到21mg/L时,再提高磷素浓度无明显增产作用；V3-R1、R1-R5期间断磷显著降低了大豆产量,R5-R7期间断磷无显著影响；在十壤速效磷含量为26.31mg·kg-1的框栽条件下,施用磷肥无明显增产作用。(3)砂培条件下,大豆产量随着钾素水平的升高而呈单峰曲线变化,64mg/L钾素浓度达到最大值,当钾素浓度达到52mg/L时,再提高钾素浓度增产作用不显著；V3-R1和R1-R5断钾显著降低了大豆产量,R5-R7断钾未达到显著差异水平；在土壤速效钾含量为174.75mg·kg-1的框栽条件下,大豆产量随钾水平的升高呈上升趋势。(4)供试大豆品种合农60、嫩丰18和绥农14的产量均随着生育期间水分的增加而提高,表明提高降水量可以达到增产的目的；大豆R4-R7对干旱最敏感,此时期干旱导致产量明显下降。
Soybean is grain and oil crop in China. Nitrogen nutrition was provided for soybean growth by soybean-rhizobium symbiosis, studying on characteristics of nodule nitrogen fixation and influencing factors-fertility and moisture conditions are profound implications for appropriate application of fertilizer, increasing yield, improving quality, reduction in production costs and alleviation of environmental pollution.
This work was conducted from2009to2011, using15N tracer technique with frame tests (pot without bottom) and sand culture. In frame experiment, DGGE and AFLP methods were used to study differences of rhizobium and nodule nitrogen fixation among different soybean varieties; effect of fertility factors and moisture conditions on nodule nitrogen fixation and yield were studied with SN14by15N tracer technique. In sand culture, effect of phosphorus nutrient and potassium nutrient on nodule nitrogen fixation and yield were researched with SN14by15N tracer technique. The results showed that:
Symbiotic rhizobium with tested soybean varieties were mainly uncultured bacterium. DGGE method was used to research differences of R4rhizobium among LX1, FS10, SN14, XJH and MSD, the number and brightness of bands were different; sequence analysis of the bands with NCBI stated that the similarity was94%-100%with uncultured bacterium from environmental samples.
AFLP results of HH41, SN14, HN40and MSD showed that there were578bands with5primer combinations, the percentage of polymorphic loci was68.86%; Rhizobium could be divided into two clusters, rhizobium of SN14R2and SN14R6were one kind, others were another kind; there was no difference of rhizobium under the condition of drought at different growth stages.
Soybean varieties with the long growth period had obvious advantages on nodule nitrogen fixation, the accumulation and ratio of nodule nitrogen fixation of tested varieties both showed that late varieties>medium varieties>early varieties, and the accumulation of nodule nitrogen fixation had significant difference.
The effect of nitrogen levels on nodule nitrogen fixation of soybean was different because of soil fertility. The accumulation and ratio of nodule nitrogen fixation of soybean presented a single peak curve with nitrogen levels when the available nitrogen content of soil was18.17mg·kg-1, nodule nitrogen fixation was the highest when the amount of (NH4)2SO4was225kg/hm2(pure N:47.73kg/hm2); nodule nitrogen fixation was the highest when the amount of (NH4)2SO4was75kg/hm2(pure N:15.91kg/hm2) under the condition of available nitrogen content of soil was 50.38mg·kg-1, nodule nitrogen fixation was significantly inhibited with the nitrogen levels increasing.
There was significant effect of phosphorus nutrient on nodule nitrogen fixation of soybean. With sand culture, the accumulation and ratio of nodule nitrogen fixation of soybean presented a single peak curve with phosphorus levels, when phosphorus concentration was11mg/L, increasing phosphorus concentration again had no obvious promotion of ratio of nodule nitrogen fixation, when phosphorus concentration was21mg/L, increasing phosphorus concentration again had no obvious promotion of accumulation of nodule nitrogen fixation; accumulation and ratio of nodule nitrogen fixation decreased significantly when interruption of phosphorus supply during V3-R1and R1-R5period, while interruption of phosphorus supply during R5-R7period had no significant effect on those, which means that nitrogen metabolism of soybean was sensitive to phosphorus requirement during V3-R5period and not sensitive after R5; with frame, phosphate application could improve the accumulation of nodule nitrogen fixation of soybean when available phosphorus content of soil was26.31mg·kg-1, while the promotion effect on accumulation of nodule nitrogen fixation of soybean was not significant with phosphate fertilizer increasing when the amount of triple superphosphate was150kg/hm2(P2O5:69kg/hm2); the ratio of nodule nitrogen fixation of soybean showed an increasing tendency with phosphate fertilizer increasing, the ratio of nodule nitrogen fixation of whole plant and seeds was significantly increased when the amount of triple superphosphate was450kg/hm2(P2O5:207kg/hm2).
There was effect of potassium nutrient on nodule nitrogen fixation of soybean. With sand culture, the accumulation and ratio of nodule nitrogen fixation of soybean presented a single peak curve with potassium levels, nodule nitrogen fixation could be satisfied when potassium levels was21mg/L; there was no significant effect of interruption of potassium supply during different periods on the accumulation and ratio of nodule nitrogen fixation; potassium fertilizer could improve the accumulation and ratio of nodule nitrogen fixation of soybean when available potassium content of soil was174.75mg·kg-1, potassium could satisfy the need of nodule nitrogen fixation when the amount of K2SO4was90kg/hm2(K2O:45kg/hm2).
The accumulation and ratio of nodule nitrogen fixation showed an increasing tendency with moisture increasing in HN60, NF18and SN14, the sensitivity of the ratio of nodule nitrogen fixation to moisture was lower than the accumulation of nodule nitrogen fixation; there was no obvious rules of nodule nitrogen fixation under the condition of phased drought during growth period.
Fertility and moisture factors could influence soybean yield.(1) Soybean yield presented a single peak curve with the increase of nitrogen rate under the condition of two soil treatments, which showed that the yield decreased either too high or too low.(2) With sand culture, soybean yield presented a single peak curve with phosphorus levels, the yield reached its maximum when phosphorus concentration was31mg/L, while increasing phosphorus concentration again had no obvious promotion of yield when phosphorus concentration was21mg/L; soybean yield decreased significantly when interruption of phosphorus supply during V3-R1and R1-R5period, while interruption of phosphorus supply during R5-R7period had no significant effect on yield; soybean yield was not significantly increased with phosphate application when available phosphorus content of soil was26.31mg·kg-1.(3) Soybean yield presented a single peak curve with potassium levels increasing with sand culture, the yield reached its maximum when potassium concentration was64mg/L, while increasing potassium concentration again had no obvious promotion of yield when potassium concentration was52mg/L; soybean yield decreased significantly when interruption of potassium supply during V3-R1and R1-R5period, while interruption of potassium supply during R5-R7period had no significant effect on yield; soybean yield showed an increasing tendency with potassium levels increasing when available potassium content of soil was174.75mg·kg-1.(4) The yield of HN60, NF18and SN14was increased with moisture improving, which showed that improving precipitation was beneficial to yield; podding-seed filling period of soybean was the most sensitive to drought, and the yield was significantly decreased.
引文
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