苦荞根系分泌有机酸对低氮胁迫的响应机制
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摘要
通过对两种不同低氮耐受性苦荞品种{迪庆苦荞(DQ,耐低氮)及黑丰1号(HF,不耐低氮)}的盆栽试验,研究了苦荞根系分泌有机酸(丙二酸、丙酸、草酸、酒石酸、乙酸)对低氮胁迫(尿素,80 mg kg~(-1))的响应机制。分析结果表明:幼苗期时,低氮处理(尿素,80 mg kg~(-1))与常氮处理(尿素,160 mg kg~(-1))相比,HF和DQ土壤含水量分别减少24.2%和14.32%,且这一时期低氮胁迫下DQ耗水量显著高于HF。DQ根际土壤pH值在幼苗期和开花期分别比HF低1.44%和8.44%。裂区分析显示,氮处理对苦荞根际土壤有机酸含量有显著的影响。低氮胁迫下苦荞根系分泌有机酸含量在品种间具有差异,根际土壤中丙二酸与其他有机酸相比含量较多但并未在品种间产生显著差异,DQ根际土壤中草酸含量在各个生育期分别显著高于HF 40.16%、25.24%和41.31%,酒石酸也在开花期和成熟期表现出相同的趋势。综上所述,苦荞可能通过根系有机酸的分泌来调节根际土壤养分有效性以应对低氮胁迫环境,对于未来贫瘠土壤上苦荞的种植应考量其耐瘠性的差异,选育耐瘠性强的品种来增加效益。
A pot experiment was conducted to study the response mechanism of organic acids(malonic acid, propionic acid, oxalic acid, tartaric acid and acetic acid) secreted from two tartary buckwheat varieties, Diqing(DQ, low nitrogen resistance) and Heifeng 1(HF, low nitrogen sensitive), to low nitrogen stress(urea, 80 mg kg~(-1)). The results showed that the soil moisture of HF and DQ under low nitrogen treatment decreased 24.2% and 14.32%, respectively when compared with normal nitrogen treatment(urea, 160 mg kg~(-1)), and the water consumption of DQ was significantly higher than that of HF at seedling stage. However, the soil pH value of DQ was 1.44% and 8.44% lower than that of HF at seedling and flowering stages, respectively. Nitrogen treatment significantly affected the contents of organic acids in buckwheat rhizosphere soil. The contents of organic acids were different among buckwheat varieties under low nitrogen stress. The content of malonic acid was higher than that of the other organic acids, but there was no significant difference among buckwheat varieties. However, the content of oxalic acid in rhizosphere soil of DQ was 40.16%, 25.24% and 41.31% higher than that of HF, respectively in each growth period, and tartaric acid also showed the same trend at flowering and maturity stage. In summary, tartary buckwheat may regulate the nutrient availability of rhizosphere soil through the secretion of organic acids in the root system to cope with the low nitrogen stress environment. The differences in the tolerance of tartary buckwheat to poor soil should be considered before its cultivation and the varieties resistant to poor soil should be bred to increase efficiency in the future.
A pot experiment was conducted to study the response mechanism of organic acids(malonic acid, propionic acid, oxalic acid, tartaric acid and acetic acid) secreted from two tartary buckwheat varieties, Diqing(DQ, low nitrogen resistance) and Heifeng 1(HF, low nitrogen sensitive), to low nitrogen stress(urea, 80 mg kg~(-1)). The results showed that the soil moisture of HF and DQ under low nitrogen treatment decreased 24.2% and 14.32%, respectively when compared with normal nitrogen treatment(urea, 160 mg kg~(-1)), and the water consumption of DQ was significantly higher than that of HF at seedling stage. However, the soil pH value of DQ was 1.44% and 8.44% lower than that of HF at seedling and flowering stages, respectively. Nitrogen treatment significantly affected the contents of organic acids in buckwheat rhizosphere soil. The contents of organic acids were different among buckwheat varieties under low nitrogen stress. The content of malonic acid was higher than that of the other organic acids, but there was no significant difference among buckwheat varieties. However, the content of oxalic acid in rhizosphere soil of DQ was 40.16%, 25.24% and 41.31% higher than that of HF, respectively in each growth period, and tartaric acid also showed the same trend at flowering and maturity stage. In summary, tartary buckwheat may regulate the nutrient availability of rhizosphere soil through the secretion of organic acids in the root system to cope with the low nitrogen stress environment. The differences in the tolerance of tartary buckwheat to poor soil should be considered before its cultivation and the varieties resistant to poor soil should be bred to increase efficiency in the future.
引文
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[3]李强,罗延宏,龙文靖,等.低氮胁迫对不同耐低氮性玉米品种苗期生长和生理特性的影响[J].草业学报,2014,23(4):204-212.
[4]张美俊,乔治军,杨武德,等.不同糜子品种对低氮胁迫的生物学响应[J].植物营养与肥料学报,2014,20(3):661-669.
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[7]路之娟,张永清,张楚,等.不同基因型苦荞苗期抗旱性综合评价及指标筛选[J].中国农业科学,2017,50(17):3311-3322.
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[11]XU H,ZHOU F,BAI K,et al.Effects of allantoin on the root growth and bleeding liquid of maize seedlings[J].Journal of Guangxi Agricultural&Biological Science,2003,22(1):25-28.
[12]张定一,张永清,杨武德,等.不同基因型小麦对低氮胁迫的生物学响应[J].小麦研究,2006,32(1):1349-1354.
[13]吴雅薇,李强,豆攀,等.低氮胁迫对不同耐低氮玉米品种苗期伤流液性状及根系活力的影响[J].植物营养与肥料学报,2017,23(2):278-288.
[14]齐泽民,卿东红.根系分泌物及其生态效应[J].内江师范学院学报,2005,20(2):68-74.
[15]赵宽,周葆华,马万征,羊礼敏.不同环境胁迫对根系分泌有机酸的影响研究进展[J].土壤,2016,48(2):235-240.
[16]SHEN H,YAN X.Exudation and Accumulation of Organic Acids in the Roots of Common Bean(Phaseolus vulgaris L.)in Response to Low Phosphorus and Aluminum Toxicity Stress[J].Acta Ecologica Sinica,2002,22(3):387-394.
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[19]张子义,伊霞,胡博,等.缺氮条件下燕麦根轴细胞的程序性死亡[J].中国农学通报,2010,26(8):175-178.
[20]DARRAH P R.Rhizodeposition under ambient and elevated CO2levels[J].Plant&Soil,1995,187(2):265-275.
[21]BAIS H P,WEIR T L,PERRY L G,et al.The role of root exudates in rhizosphere interactions with plants and other organisms[J].Annual Review of Plant Biology,2006,57(1):233-266.
[22]董艳,董坤,郑毅,等.不同品种小麦与蚕豆间作对蚕豆枯萎病的防治及其机理[J].应用生态学报,2014,25(7):1979-1987.
[23]CHEN W,ZHANG L L,LI X Y,et al.Elevated ozone increases nitrifying and denitrifying enzyme activities in the rhizosphere of wheat after 5 years of fumigation.Plant&Soil,2015,392(1-2):279-288.
[24]李德华,向春雷,姜益泉,等.低磷胁迫下水稻不同品种根系有机酸分泌的差异[J].中国农学通报,2005,21(11):186-188.
[25]孙宝利,赤杰,范中南,等.土壤及植物复合体系中有机酸的测定[J].环境科学与技术,2010,33(9):137-141.
[26]李煜姗,杨再强,李平,等.高效液相色谱法测定设施番茄土壤低分子量有机酸的色谱条件研究[J].土壤通报,2016,47(1):73-78.
[27]陈伟,皇甫倩华,孙从建,等.大气O3升高对小麦根际土壤微生物量和氮素化酶活性的影响[J].土壤通报,2017,48(3):623-630.
[28]陈龙池,廖利平,汪思龙,等.根系分泌物生态学研究[J].生态学杂志,2002,21(6):57-62.
[29]徐国伟,李帅,赵永芳,等.秸秆还田与施氮对水稻根系分泌物及氮素利用的影响研究[J].草业学报,2014,23(2):140-146.
[30]杨迪,马瑞,黄文斌,等.养分和水分胁迫下2年生落叶松根系有机酸的分泌行为研究[J].安徽农业科学,2013(36):13932-13934.
[31]汪建飞,沈其荣.有机酸代谢在植物适应养分和铝毒胁迫中的作用[J].应用生态学报,2006,17(11):2210-2216.
[32]张俊英,王敬国,许永利,等.氮素对不同大豆品种根系分泌物中有机酸的影响[J].植物营养与肥料学报,2007,13(3):398-403.
[33]赵宽,周葆华,马万征,羊礼敏.不同环境胁迫对根系分泌有机酸的影响研究进展[J].土壤,2016,48(2):235-240.
[34]李振侠,徐继忠,高仪,等.苹果砧木SH40和八棱海棠缺铁胁迫下根系有机酸分泌的差异[J].园艺学报,2007,34(2):279-282.
[35]DINKELAKER B,R魻MHELD V,MARSCHNER H.Citric acid excretion and precipitation of calcium citrate in the rhizosphere of white lupin(Lupinus albus L.).Plant Cell&Environment,1989,12(3):285-292.
[36]王妍.养分缺乏下有机酸对暗棕壤铁活化和落叶松幼苗吸收铁的影响[J].林业科技情报,2012,(4):18-21.
[37]常二华,张耗,张慎凤,等.结实期氮磷营养水平对水稻根系分泌物的影响及其与稻米品质的关系[J].作物学报,2007,33(12):1949-1959.
[38]戢林,李廷轩,张锡洲,等.水稻氮高效基因型根系分泌物中有机酸和氨基酸的变化特征[J].植物营养与肥料学报,2012,18(5):1046-1055.
[39]罗燕,樊卫国.不同施磷水平下4种柑橘砧木的根际土壤有机酸、微生物及酶活性[J].中国农业科学,2014,47(5):955-967.
[2]王业建.大豆对低氮胁迫的形态和生理学响应及介导低氮胁迫mi RNA的鉴定[D].中南大学,2013.
[3]李强,罗延宏,龙文靖,等.低氮胁迫对不同耐低氮性玉米品种苗期生长和生理特性的影响[J].草业学报,2014,23(4):204-212.
[4]张美俊,乔治军,杨武德,等.不同糜子品种对低氮胁迫的生物学响应[J].植物营养与肥料学报,2014,20(3):661-669.
[5]张楚,张永清,路之娟,等.低氮胁迫对不同苦荞品种苗期生长和根系生理特征的影响[J].西北植物学报,2017,37(7):1331-1339.
[6]倪志巍.山西省小杂粮产业现状与发展对策[D].山西农业大学,2016.
[7]路之娟,张永清,张楚,等.不同基因型苦荞苗期抗旱性综合评价及指标筛选[J].中国农业科学,2017,50(17):3311-3322.
[8]张楚,张永清,路之娟,等.苗期耐低氮基因型苦荞的筛选及其评价指标[J].作物学报,2017,43(8):1205-1215.
[9]SEITH B,GEORGE E,MARSCHNER H,et al.Effects of varied soil nitrogen supply on Norway spruce(Picea abies,[L.]Karst.)[J].Plant&Soil,1996,184(2):291-298.
[10]谢孟林,李强,查丽,等.低氮胁迫对不同耐低氮性玉米品种幼苗根系形态和生理特征的影响[J].中国生态农业学报,2015,23(8):946-953.
[11]XU H,ZHOU F,BAI K,et al.Effects of allantoin on the root growth and bleeding liquid of maize seedlings[J].Journal of Guangxi Agricultural&Biological Science,2003,22(1):25-28.
[12]张定一,张永清,杨武德,等.不同基因型小麦对低氮胁迫的生物学响应[J].小麦研究,2006,32(1):1349-1354.
[13]吴雅薇,李强,豆攀,等.低氮胁迫对不同耐低氮玉米品种苗期伤流液性状及根系活力的影响[J].植物营养与肥料学报,2017,23(2):278-288.
[14]齐泽民,卿东红.根系分泌物及其生态效应[J].内江师范学院学报,2005,20(2):68-74.
[15]赵宽,周葆华,马万征,羊礼敏.不同环境胁迫对根系分泌有机酸的影响研究进展[J].土壤,2016,48(2):235-240.
[16]SHEN H,YAN X.Exudation and Accumulation of Organic Acids in the Roots of Common Bean(Phaseolus vulgaris L.)in Response to Low Phosphorus and Aluminum Toxicity Stress[J].Acta Ecologica Sinica,2002,22(3):387-394.
[17]HOFFLAND E,VAN DEN BOOGAARD R,NELEMANS J,et al.Biosynthesis and root exudation of citric and malic acid in phosphate-starved rape plants[J].New Phytologist,1992,122(4):675-680.
[18]KONINGS H,VERSCHUREN G.Formation of aerenchyma in roots of Zea mays in aerated solutions,and its relation to nutrient supply[J].Physiol.Plant.,1980,49:265-279.
[19]张子义,伊霞,胡博,等.缺氮条件下燕麦根轴细胞的程序性死亡[J].中国农学通报,2010,26(8):175-178.
[20]DARRAH P R.Rhizodeposition under ambient and elevated CO2levels[J].Plant&Soil,1995,187(2):265-275.
[21]BAIS H P,WEIR T L,PERRY L G,et al.The role of root exudates in rhizosphere interactions with plants and other organisms[J].Annual Review of Plant Biology,2006,57(1):233-266.
[22]董艳,董坤,郑毅,等.不同品种小麦与蚕豆间作对蚕豆枯萎病的防治及其机理[J].应用生态学报,2014,25(7):1979-1987.
[23]CHEN W,ZHANG L L,LI X Y,et al.Elevated ozone increases nitrifying and denitrifying enzyme activities in the rhizosphere of wheat after 5 years of fumigation.Plant&Soil,2015,392(1-2):279-288.
[24]李德华,向春雷,姜益泉,等.低磷胁迫下水稻不同品种根系有机酸分泌的差异[J].中国农学通报,2005,21(11):186-188.
[25]孙宝利,赤杰,范中南,等.土壤及植物复合体系中有机酸的测定[J].环境科学与技术,2010,33(9):137-141.
[26]李煜姗,杨再强,李平,等.高效液相色谱法测定设施番茄土壤低分子量有机酸的色谱条件研究[J].土壤通报,2016,47(1):73-78.
[27]陈伟,皇甫倩华,孙从建,等.大气O3升高对小麦根际土壤微生物量和氮素化酶活性的影响[J].土壤通报,2017,48(3):623-630.
[28]陈龙池,廖利平,汪思龙,等.根系分泌物生态学研究[J].生态学杂志,2002,21(6):57-62.
[29]徐国伟,李帅,赵永芳,等.秸秆还田与施氮对水稻根系分泌物及氮素利用的影响研究[J].草业学报,2014,23(2):140-146.
[30]杨迪,马瑞,黄文斌,等.养分和水分胁迫下2年生落叶松根系有机酸的分泌行为研究[J].安徽农业科学,2013(36):13932-13934.
[31]汪建飞,沈其荣.有机酸代谢在植物适应养分和铝毒胁迫中的作用[J].应用生态学报,2006,17(11):2210-2216.
[32]张俊英,王敬国,许永利,等.氮素对不同大豆品种根系分泌物中有机酸的影响[J].植物营养与肥料学报,2007,13(3):398-403.
[33]赵宽,周葆华,马万征,羊礼敏.不同环境胁迫对根系分泌有机酸的影响研究进展[J].土壤,2016,48(2):235-240.
[34]李振侠,徐继忠,高仪,等.苹果砧木SH40和八棱海棠缺铁胁迫下根系有机酸分泌的差异[J].园艺学报,2007,34(2):279-282.
[35]DINKELAKER B,R魻MHELD V,MARSCHNER H.Citric acid excretion and precipitation of calcium citrate in the rhizosphere of white lupin(Lupinus albus L.).Plant Cell&Environment,1989,12(3):285-292.
[36]王妍.养分缺乏下有机酸对暗棕壤铁活化和落叶松幼苗吸收铁的影响[J].林业科技情报,2012,(4):18-21.
[37]常二华,张耗,张慎凤,等.结实期氮磷营养水平对水稻根系分泌物的影响及其与稻米品质的关系[J].作物学报,2007,33(12):1949-1959.
[38]戢林,李廷轩,张锡洲,等.水稻氮高效基因型根系分泌物中有机酸和氨基酸的变化特征[J].植物营养与肥料学报,2012,18(5):1046-1055.
[39]罗燕,樊卫国.不同施磷水平下4种柑橘砧木的根际土壤有机酸、微生物及酶活性[J].中国农业科学,2014,47(5):955-967.