丛枝菌根真菌提高黄土高原紫穗槐抗旱能力的研究
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摘要
本文在陕西黄土高原的甘泉、绥德、米脂、榆林4县（市）选取4个样地，研究紫穗槐（Amorpha fruticosa）不同深度土层丛枝菌根真菌（Arbuscular mycorrhizal fungi，AMF）侵染率、孢子密度、球囊霉素和土壤因子之间的关系，并利用盆栽试验研究了水分胁迫下接种丛枝菌根真菌对紫穗槐生长和抗旱性的影响。得出以下主要结论：
1. AMF侵染率平均值绥德样地最大，为100%，米脂样地最小，为75.02%；孢子密度平均值米脂样地最大，为5.91个/g，榆林样地最小，为1.57个/g。4个样地孢子密度最大值均在0~(-1)0cm土层，并且随土层加深而降低。AMF侵染率与孢子密度在各样地间差异显著，同一样地侵染率与孢子密度变化规律不一致。榆林样地的有机碳、铵态氮、速效磷、脲酶和碱性磷酸酶活性平均值显著高于其他样地。除绥德、米脂样地的蔗糖酶和甘泉样地的碱性磷酸酶外，其他样地的土壤酶活性和球囊霉素含量均随土层加深而降低，且各土层之间差异显著，最大值均在0~(-1)0cm土层。
2. AMF侵染率与土壤pH、总球囊霉素和易提取球囊霉素含量呈极显著正相关，与有机碳、硝态氮、速效磷含量呈显著正相关。孢子密度与速效磷含量、蔗糖酶和脲酶活性呈显著正相关，与铵态氮含量呈显著负相关。主成分分析表明，第1、2主成分的累积方差贡献率达到56.4%，第1主成分主要综合了球囊霉素含量、AMF侵染率和孢子密度；第2主成分综合了土壤pH和部分土壤因子信息。因此，球囊霉素含量、AMF侵染率、孢子密度、pH、部分土壤因子对决定土壤生态起主要作用。
3.水分胁迫下（土壤相对含水量为30%），接种植株的根系活力（0.66mg g~(-1) h~(-1)）显著高于对照（0.49mg g~(-1) h~(-1)）；随着土壤相对含水量从70%降至30%，接种和对照的植株株高、地上和地下部干重、根系活力均下降，而接种的菌根侵染率则从43.99%提高到88.47%，显著上升；接种丛枝菌根真菌显著增加了叶片脯氨酸、可溶性糖和可溶性蛋白含量，降低了叶片丙二醛含量，尤其到胁迫后期，土壤相对含水量为30%时接种植株的脯氨酸含量为135.67μg/g，为对照（58.72μg/g）的2.31倍。土壤相对含水量为50%时接种植株SOD、POD、CAT的活性分别是对照的1.47、1.48和1.37倍。植株在长时间受到水分胁迫时，接种AMF可以显著提高这三种酶的活性。
4.水分胁迫下，接种植株的Fv/Fm显著高于对照。土壤相对含水量为30%的对照，随着水分胁迫时间的延长，Fv/Fm在中后期从0.76快速降低至0.70，而接种则始终维持在0.78左右。水分胁迫前期各水分处理ETR值差异不显著，随着水分胁迫时间的延续，ETR值呈先降低再升高的趋势，土壤相对含水量为30%时的对照在中期下降幅度最大，从9.8降至6.0，且后期不能升高到原有水平，而接种可以恢复到接近原水平。随着土壤相对含水量的增加，qL显著上升，接种高于对照。水分胁迫至中期，土壤相对含水量为30%的接种和对照qL显著下降，从0.86降至0.73。胁迫至后期，对照qL继续降低至0.60，而接种回升至0.84，接近前期水平。各处理在水分胁迫初期，NPQ随土壤相对含水量的降低而上升，水分胁迫延续至中后期，各处理NPQ均缓慢上升，对照高于接种。
We collected the roots and rhizosphere soil samples from four different sites (Ganquan, Suide, Mizhiand Yulin) on the Loess Plateau in Shaanxi. The spatial distribution of AMF colonization, spore density andglomalin in the rhizosphere of Amorpha fruticosa were investigated, and their relationships with soil factorswere analysed. Then, under pot condition, an experiment was conducted to study the effects of AMF on thegrowth and drought resistance of Amorpha fruticosa under water stress. The main conclusions included asfollows:
1. The average AMF colonization rate from Suide was the highest of all the samples, which was up to100%, and the lowest colonization rate was75.02%from Mizhi. The spore density from Mizhi was higherthan that of other samples, and it was5.91/g. The lowest spore density was from Yulin, and it was1.57/g.The highest spore density existed in the0~(-1)0cm layer of all samples, and with the soil depth increasing, thespore density decreased. In different regions, the difference of AMF colonization rate and spore densitywere notable. The rate of AMF colonization did not significantly correlate with the spore density. SOC,ammonia-N, olsen-P, urease and alkaline phosphatase from Yulin were significantly higher than those ofother samples. Except for the sucrase from Mizhi and Suide and alkaline phosphatase from Ganquan, in0~(-1)0cm layer, the activity of most soil enzymes and glomalin reached the maximum, and declined with theincreasing soil depth, which had a significant difference among the different layers.
2. AMF colonization had a significant positive correlation with SOC, nitrate-N, olsen-P, and it had avery significant positive correlation with pH, TEG and EEG. Spore density had a significant positivecorrelation with olsen-P, sucrase, urease, and had a negative correlation with ammonia-N. TEG had a verysignificant positive correlation with EEG. Principal component analysis showed that the cumulativevariance proportion of PCA1and PCA2were up to56.4%. PCA1contained glomalin, AMF colonizationrate and spore density. PCA2contained pH and some soil factors. Therefore, the soil environment wasmainly decided by these factors.
3. Under30%of soil relative water content, the root activity of Amorpha fruticosa inoculating AMFwhich was reached up to0.66mg g~(-1) h~(-1) was significantly higher than that without inoculating AMFwhich was0.49mg g~(-1) h~(-1). With soil relative water content decreasing, plant height, root activity, shootdry mass and root dry mass were decreased, and AMF colonization rate was significantly increased from43.99%to88.47%. Under the different soil relative water content, contents of proline, soluble sugar andsoluble protein were significantly increased by inoculating AMF, especially the content of proline thatinoculating AMF was2.31times as much as that without inoculating AMF, and the content of MDA wasdecrease with soil relative water content increased. Inoclulating AMF could promote the activities of threekinds of anti-oxidative enzymes included SOD, POD and CAT, which were1.47,1.48and1.37times that ofthose without inoculating AMF, and keep them at a high level.
4. Under water stress, Fv/Fm of the plant inoculating AMF was significantly higher than thatwithout inoculating AMF. Under30%of soil relative water content, Fv/Fm of the plant without inoculatingquickly decreased from0.76to0.70, when the water stress reached to the middle and later period. But frombeginning to the end, Fv/Fm with inoculating AMF was at the level about0.78. At the early period of waterstress, ETR was at the same level among the different treatment. With the days lasting, ETR was higher thanthe level of early period. Under30%of soil relative water content, the plant without inoculating AMFdecreased by a largest margin which was from9.8to6.0in middle period, and it could not get recover tothe original level at the later period. But the plant inoculating AMF could get recover. qL increasedsignificantly with the increasing of soil relative water content, and the plant inoculating AMF was higherthan that without inoculating AMF. At the middle of water stress, qL decreased significantly from0.86to0.73under30%of soil relative water content. The plant without inoculating AMF continueded to drop to0.6,but the plant inoculating AMF get recover to original level about0.84. At the early period of waterstress, NPQ increased with the decreasing of soil relative water content. With the days lasting, NPQ of alltreatments increased slowly, and the plant of inoculating AMF was lower than that without inoculatingAMF.
1. AMF侵染率平均值绥德样地最大，为100%，米脂样地最小，为75.02%；孢子密度平均值米脂样地最大，为5.91个/g，榆林样地最小，为1.57个/g。4个样地孢子密度最大值均在0~(-1)0cm土层，并且随土层加深而降低。AMF侵染率与孢子密度在各样地间差异显著，同一样地侵染率与孢子密度变化规律不一致。榆林样地的有机碳、铵态氮、速效磷、脲酶和碱性磷酸酶活性平均值显著高于其他样地。除绥德、米脂样地的蔗糖酶和甘泉样地的碱性磷酸酶外，其他样地的土壤酶活性和球囊霉素含量均随土层加深而降低，且各土层之间差异显著，最大值均在0~(-1)0cm土层。
2. AMF侵染率与土壤pH、总球囊霉素和易提取球囊霉素含量呈极显著正相关，与有机碳、硝态氮、速效磷含量呈显著正相关。孢子密度与速效磷含量、蔗糖酶和脲酶活性呈显著正相关，与铵态氮含量呈显著负相关。主成分分析表明，第1、2主成分的累积方差贡献率达到56.4%，第1主成分主要综合了球囊霉素含量、AMF侵染率和孢子密度；第2主成分综合了土壤pH和部分土壤因子信息。因此，球囊霉素含量、AMF侵染率、孢子密度、pH、部分土壤因子对决定土壤生态起主要作用。
3.水分胁迫下（土壤相对含水量为30%），接种植株的根系活力（0.66mg g~(-1) h~(-1)）显著高于对照（0.49mg g~(-1) h~(-1)）；随着土壤相对含水量从70%降至30%，接种和对照的植株株高、地上和地下部干重、根系活力均下降，而接种的菌根侵染率则从43.99%提高到88.47%，显著上升；接种丛枝菌根真菌显著增加了叶片脯氨酸、可溶性糖和可溶性蛋白含量，降低了叶片丙二醛含量，尤其到胁迫后期，土壤相对含水量为30%时接种植株的脯氨酸含量为135.67μg/g，为对照（58.72μg/g）的2.31倍。土壤相对含水量为50%时接种植株SOD、POD、CAT的活性分别是对照的1.47、1.48和1.37倍。植株在长时间受到水分胁迫时，接种AMF可以显著提高这三种酶的活性。
4.水分胁迫下，接种植株的Fv/Fm显著高于对照。土壤相对含水量为30%的对照，随着水分胁迫时间的延长，Fv/Fm在中后期从0.76快速降低至0.70，而接种则始终维持在0.78左右。水分胁迫前期各水分处理ETR值差异不显著，随着水分胁迫时间的延续，ETR值呈先降低再升高的趋势，土壤相对含水量为30%时的对照在中期下降幅度最大，从9.8降至6.0，且后期不能升高到原有水平，而接种可以恢复到接近原水平。随着土壤相对含水量的增加，qL显著上升，接种高于对照。水分胁迫至中期，土壤相对含水量为30%的接种和对照qL显著下降，从0.86降至0.73。胁迫至后期，对照qL继续降低至0.60，而接种回升至0.84，接近前期水平。各处理在水分胁迫初期，NPQ随土壤相对含水量的降低而上升，水分胁迫延续至中后期，各处理NPQ均缓慢上升，对照高于接种。
We collected the roots and rhizosphere soil samples from four different sites (Ganquan, Suide, Mizhiand Yulin) on the Loess Plateau in Shaanxi. The spatial distribution of AMF colonization, spore density andglomalin in the rhizosphere of Amorpha fruticosa were investigated, and their relationships with soil factorswere analysed. Then, under pot condition, an experiment was conducted to study the effects of AMF on thegrowth and drought resistance of Amorpha fruticosa under water stress. The main conclusions included asfollows:
1. The average AMF colonization rate from Suide was the highest of all the samples, which was up to100%, and the lowest colonization rate was75.02%from Mizhi. The spore density from Mizhi was higherthan that of other samples, and it was5.91/g. The lowest spore density was from Yulin, and it was1.57/g.The highest spore density existed in the0~(-1)0cm layer of all samples, and with the soil depth increasing, thespore density decreased. In different regions, the difference of AMF colonization rate and spore densitywere notable. The rate of AMF colonization did not significantly correlate with the spore density. SOC,ammonia-N, olsen-P, urease and alkaline phosphatase from Yulin were significantly higher than those ofother samples. Except for the sucrase from Mizhi and Suide and alkaline phosphatase from Ganquan, in0~(-1)0cm layer, the activity of most soil enzymes and glomalin reached the maximum, and declined with theincreasing soil depth, which had a significant difference among the different layers.
2. AMF colonization had a significant positive correlation with SOC, nitrate-N, olsen-P, and it had avery significant positive correlation with pH, TEG and EEG. Spore density had a significant positivecorrelation with olsen-P, sucrase, urease, and had a negative correlation with ammonia-N. TEG had a verysignificant positive correlation with EEG. Principal component analysis showed that the cumulativevariance proportion of PCA1and PCA2were up to56.4%. PCA1contained glomalin, AMF colonizationrate and spore density. PCA2contained pH and some soil factors. Therefore, the soil environment wasmainly decided by these factors.
3. Under30%of soil relative water content, the root activity of Amorpha fruticosa inoculating AMFwhich was reached up to0.66mg g~(-1) h~(-1) was significantly higher than that without inoculating AMFwhich was0.49mg g~(-1) h~(-1). With soil relative water content decreasing, plant height, root activity, shootdry mass and root dry mass were decreased, and AMF colonization rate was significantly increased from43.99%to88.47%. Under the different soil relative water content, contents of proline, soluble sugar andsoluble protein were significantly increased by inoculating AMF, especially the content of proline thatinoculating AMF was2.31times as much as that without inoculating AMF, and the content of MDA wasdecrease with soil relative water content increased. Inoclulating AMF could promote the activities of threekinds of anti-oxidative enzymes included SOD, POD and CAT, which were1.47,1.48and1.37times that ofthose without inoculating AMF, and keep them at a high level.
4. Under water stress, Fv/Fm of the plant inoculating AMF was significantly higher than thatwithout inoculating AMF. Under30%of soil relative water content, Fv/Fm of the plant without inoculatingquickly decreased from0.76to0.70, when the water stress reached to the middle and later period. But frombeginning to the end, Fv/Fm with inoculating AMF was at the level about0.78. At the early period of waterstress, ETR was at the same level among the different treatment. With the days lasting, ETR was higher thanthe level of early period. Under30%of soil relative water content, the plant without inoculating AMFdecreased by a largest margin which was from9.8to6.0in middle period, and it could not get recover tothe original level at the later period. But the plant inoculating AMF could get recover. qL increasedsignificantly with the increasing of soil relative water content, and the plant inoculating AMF was higherthan that without inoculating AMF. At the middle of water stress, qL decreased significantly from0.86to0.73under30%of soil relative water content. The plant without inoculating AMF continueded to drop to0.6,but the plant inoculating AMF get recover to original level about0.84. At the early period of waterstress, NPQ increased with the decreasing of soil relative water content. With the days lasting, NPQ of alltreatments increased slowly, and the plant of inoculating AMF was lower than that without inoculatingAMF.
引文
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