不同供氮形态下水稻苗期磷吸收累积与根系形态的关系
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摘要
【目的】植物根系形态对于适应低磷胁迫具有一定的可塑性,对提高磷的吸收利用具有重要意义。因此,本研究以长江中下游地区主推的102个水稻品种为供试材料,研究根系形态与水稻幼苗磷吸收利用的相关性。【方法】采用国际水稻所营养液培养方法,研究在NH_4~+-N和NO_3~--N供应条件下苗期植株生物量、磷含量和磷素累积量及其与根系形态指标的相关性。【结果】研究结果表明,在相同供氮水平(40 mg/L)下,供应NH_4~+-N时,水稻苗期平均生物量为67.87 mg/株,比供应NO_3~--N时高4.27 mg/株;水稻苗期平均磷含量为0.49%,比供应NO_3~--N时高0.10%;水稻苗期平均磷累积量为0.37 mg/株,比供应NO_3~--N时高0.10 mg/株。在NH_4~+-N条件下,水稻根系形态指标变异系数呈现根尖数>总根长>分支数>总根面积>交叉数>总根体积>平均根系直径的规律;在NO_3~--N条件下,水稻根系形态指标变异系数呈现根尖数>分枝数>总根长>总根面积>交叉数>总根体积>平均根系直径的趋势。在NH_4~+-N条件下,总根长、总根面积、分枝数、交叉数四个形态指标与植株生物量、磷含量、磷累积相关最为显著(P<0.01),而在NO_3~--N培养下,总根长、总根面积、根尖数、交叉数与植株生物量及磷素吸收累积指标相关性最为显著(P<0.01)。【结论】供应氨态氮,水稻营养指标与根系形态指标的相关性更高。水稻苗期根系总根长、总根面积、交叉数可作为水稻磷高效评价的重要指标。
【Objective】Plant root morphology has certain plasticity for adapting to low phosphorus stress, and is of great significance for improving phosphorus absorption and utilization. Therefore, we used 102 major rice varieties in the middle and lower reaches of the Yangtze River as the tested materials to study the correlation between root morphology and phosphorus uptake of rice seedlings. 【Method】Hydroponic experiments were conducted to investigate the influences of NH_4~+-N and NO_3~--N at the same concentration(40 mg·L-1) on dry weight, P concentrations and contents as well as the root morphology.【Result】At the six-leaf stage of rice incubated under the same concentration of nitrogen, the average biomass of the rice seedlings amounted to 67.87 mg per plant in the NH_4~+-N solution, 4.27 mg higher than that in the NO_3~--N solution. The average P content was 0.49% when grown in the NH_4~+-N solution, 0.10% higher than that of rice seedlings grown in the NO_3~--N solution. In addition, the average accumulation of P in the seedlings was 0.37 mg per plant when supplied with NH_4~+-N, 0.10 mg higher than that supplied with NO_3~--N. The coefficient of variation of root morphological indexes were in the order of the number of tips > total root length > the number of branches > total root area > the number of crossing > the total root volume >average diameter in the NH_4~+-N solution; the coefficient of variation of root morphological indexes were in the order of the number of tips > the number of branches > total root length > total root area > the number of crossing >the total root volume > average diameter in the NO_3~--N solution. The total root length, total root area, branching number and crossing number were significantly correlated with biomass, P content, and P accumulation in rice plants in the NH_4~+-N solution; whereas the total root length, total root area, tip number and crossing number were significantly correlated with biomass, P content, and P accumulation in rice plants in the NO_3~--N solution.【Conclusion】The correlations between P uptake and the indices of root morphology when plants were supplied with NH_4~+-N were higher than those supplied only with NO_3~--N. The total root length, total root area and crossing number might be important indicators for evaluating the P use efficiency of rice at seedling stage.
【Objective】Plant root morphology has certain plasticity for adapting to low phosphorus stress, and is of great significance for improving phosphorus absorption and utilization. Therefore, we used 102 major rice varieties in the middle and lower reaches of the Yangtze River as the tested materials to study the correlation between root morphology and phosphorus uptake of rice seedlings. 【Method】Hydroponic experiments were conducted to investigate the influences of NH_4~+-N and NO_3~--N at the same concentration(40 mg·L-1) on dry weight, P concentrations and contents as well as the root morphology.【Result】At the six-leaf stage of rice incubated under the same concentration of nitrogen, the average biomass of the rice seedlings amounted to 67.87 mg per plant in the NH_4~+-N solution, 4.27 mg higher than that in the NO_3~--N solution. The average P content was 0.49% when grown in the NH_4~+-N solution, 0.10% higher than that of rice seedlings grown in the NO_3~--N solution. In addition, the average accumulation of P in the seedlings was 0.37 mg per plant when supplied with NH_4~+-N, 0.10 mg higher than that supplied with NO_3~--N. The coefficient of variation of root morphological indexes were in the order of the number of tips > total root length > the number of branches > total root area > the number of crossing > the total root volume >average diameter in the NH_4~+-N solution; the coefficient of variation of root morphological indexes were in the order of the number of tips > the number of branches > total root length > total root area > the number of crossing >the total root volume > average diameter in the NO_3~--N solution. The total root length, total root area, branching number and crossing number were significantly correlated with biomass, P content, and P accumulation in rice plants in the NH_4~+-N solution; whereas the total root length, total root area, tip number and crossing number were significantly correlated with biomass, P content, and P accumulation in rice plants in the NO_3~--N solution.【Conclusion】The correlations between P uptake and the indices of root morphology when plants were supplied with NH_4~+-N were higher than those supplied only with NO_3~--N. The total root length, total root area and crossing number might be important indicators for evaluating the P use efficiency of rice at seedling stage.
引文
[1]Liedgens M,Soldati A,Stamp P.Root development of maize(Zea mays L.)as observed with minirhizotrons in lysimeters.Crop Sci,2000,40(6):1665-1672.
[2]Yan X,Liao H,Beebe S E,Blair M W,Lynch J P.QTLmapping of root hair and acid exudation traits and their relationship to phosphorus uptake in common bean.Plant&Soil,2004,265(1/2):17-29.
[3]Bates T R,Lynch J P.Stimulation of root hair elongation in Arabidopsis thaliana by low phosphorus availability.Plant Cell&Environ,1996,19(5):529-538.
[4]Jungk A.Root hairs and the acquisition of plant nutrients from soil.J Plant Nutr Soil Sci,2015,164(2):121-129
[5]Bonser A M,Lynch J,Snapp S.Effect of phosphorus deficiency on growth angle of basal roots in phaseolus Vulgaris.New Phytol,1996,132(2):281-288.
[6]Liao H,Rubio G,Yan X,Cao A Q,Brown K M,Lynch J P.Effect of phosphorus availability on basal root shallowness in common bean.Plant&Soil,2001,232(1/2):69-79
[7]沈宏,严小龙.根系分泌作用及其诱导机制.生态环境学报,2001,10(4):339-342.Sheng H,Yan X L.Root exudation and their induced mechanism.Ecol Environ Sci,2001,10(4):339-342(in Chinese with English abstract)
[8]张平.氮形态对低磷胁迫下苗期玉米磷素吸收的影响及其机理.南京:南京农业大学,2012.Zhang P.Mechanisms and effects of nitrogen forms on phosphorus absorption of maize(Zea mays L.)under phosphorus stress at seedling stage.Nanjing:Nanjing Agricultural University,2012.(in Chinese with English abstract)
[9]邹春琴,杨志福.氮素形态对春小麦根际pH与磷素营养状况的影响.土壤通报,1994(4):175-177.Zhou C Q,Yang Z F.Effects of different form nitrogen on pH and phosphorus content of spring wheat rhizosphere.Chin J Soil Sci,1994(4):175-177.(in Chinese with English abstract)
[10]李朝霞.磷匮乏影响玉米根系发育机制的研究.济南:山东大学,2011.Li C X.Study on the mechanism of root developmental changes to phosphate deficiency in maize.Jinan:Shandong University,2011.(in Chinese with English abstract)
[11]Chen X P,Cui Z L,Vitousek P M,Cassman K G,Matson P A,Bai J S.Integrated soil-crop system management for food security.PNAS,2011,108(16):6399-6404.
[12]严小龙,廖红,戈振扬.植物根构型特性与磷吸收效率.植物学报,2000,17(6):511-519.Yan X L,Liao H,Ge Z Y.root architectural characteristics and phosphorus acquisition efficiency in plants.Chin Bull Bot,2000,17(6):511-519.(in Chinese)
[13]赵静,付家兵,廖红,罗锡文.大豆磷效率应用核心种质的根构型性状评价.科学通报,2004,49(13):1249-1257.Zhao J,Fu J B,Liao H,Luo X W.Properties evaluation of phosphorus efficiency on roots structure of soybean.Chin Sci Bull,2004,49(13):1249-1257.(in Chinese)
[14]Forde B,Lorenzo H.The nutritional control of root development.Interactions in the root environment:An Integrated Approach.2002:51-68.
[15]刘灵,廖红,王秀荣,严小龙.不同根构型大豆对低磷的适应性变化及其与磷效率的关系.中国农业科学,2008,41(4):1089-1099.Liu L,Liao H,Wang X R,Yan X L.Adaptive changes of soybean genotypes with different root architectures to low phosphorus availability as related to phosphorus efficiency.Sci Agric Sin,2008,41(4):1089-1099.(in Chinese with English abstract)
[16]李佳.水稻根系应答根际水分与磷素环境变化的研究.扬州:扬州大学,2015.Li J.Study on rhizosphere moisture and phosphorus environment change of rice root response.Yangzhou:Yangzhou University,2015.(in Chinese with English abstract)
[17]陈晨,龚海青,张敬智,郜红建.水稻根系形态与氮素吸收累积的相关性分析.植物营养与肥料学报,2017,23(2):333-341.Chen C,Gong H Q,Zhang J Z,Gao H J.Correlation between root morphology and nitrogen uptake of rice.J Plant Nutr Fert,2017,23(2):333-341.(in Chinese with English abstract)
[18]Lynch J.Root Architecture and Plant Productivity.Plant Physiol,1995,109(1):7-13.
[19]李宝珍,王松伟,冯慧敏,徐国华.氮素供应形态对水稻根系形态和磷吸收的影响.中国水稻科学,2008,22(6):665-668.Li B Z,Wang S W,Feng H M,Xu G H.Effects of nitrogen forms on root morphology and phosphate uptake in rice.Chin J Rice Sci,2008,22(6):665-668(in Chinese with English abstract)
[20]Ladha J K,Kirk G J,Bennett J,Peng S,Reddy C K,Reddy P M,Singh U.Opportunities for increased nitrogen-use efficiency from improved lowland rice germplasm.Field Crops Res,1998,56(1/2):41-71.
[21]廖红,戈振扬,戈振场,严小龙.水磷耦合胁迫下植物磷吸收的理想根构型:模拟与应用.科学通报,2001,46(8):641-646.Liao H,Ge Z Y,Yan X L,Yan X L.Optimized root architecture for phosphorus acquisition under water and phosphorus stress:Simulation and application.Chin Sci Bull,2001,46(8):641-646.(in Chinese with English abstract)
[22]Muchhal U S,Pardo J M,Raghothama K G.Phosphate transporters from the higher plant Arabidopsis thaliana.PNAS,1996,93(19):10519.
[23]李宝珍,辛伟杰,徐国华.氮饥饿水稻利用不同形态氮素的差异及其生理机制.土壤学报,2007,44(2):273-279.Li B Z,Xin W J,Xu G H.Physiological mechanisms in uptake and use of different forms of nitrogen by nitrogen starved rice crop.Acta Pedol Sin,2007,44(2):273-279.(in Chinese with English abstract)
[24]范志强.氮磷营养及氮形态对水曲柳幼苗生长和生理的影响机制.沈阳:东北林业大学,2004.Fan Z Q.Effects of nitrogen,phosphorus and nitrogen forms on growth and physiological traits of fraxinus mandshurica seedlings.Shenyang:Northeast Forestry University,2004(in Chinese with English abstract)
[25]张彦东,范志强,王庆成,王政权.不同形态N素对水曲柳幼苗生长的影响.应用生态学报,2000,11(5):665-667.Zhou Y D,Fan Z Q,Wang Q C,Wang Z Q.Effects of different nitrogen forms on growth of Fraxinus mandshurica seedlings.Chin J Appl Ecol,2000,11(5):665-667.(in Chinese with English abstract)
[26]Miller M H.Effects of nitrogen on phosphorus absorption by plants.Proc Plant Root Environ,1974.
[27]Thien S J,Mcfee W W.Effect of Nitrogen on phosphorus transport systems in Zea Mays L.Soil Sci Soc Amer J,1972,36(4):617-620.
[28]Smith F W,Jackson W A.Nitrogen enhancement of phosphate transport in roots of Zea mays L.:I.Effects of ammonium and nitrate pretreatment.Plant Physiol,1987,84(4):1314-1318.
[29]Hirel B,Limami A M.Prospects for improving nitrogen use efficiency:Insights given by 15N-labelling experiments.Phytochem Rev,2002,2(1/2):133-144.
[30]Radin J W.Differential regulation of nitrate reductase induction in roots and shoots of cotton plants.Plant Physiol,1975,55(2):178-182.
[31]Zhang H,Xue YG,Wang ZQ,Yang J,Zhang J.Morphological and physiological traits of roots and their relationships with shoot growth in“super”rice.Field Crops Res,2009,113(1):31-40.
[32]Mackown C T,Jackson W A,Volk R J.Restricted nitrate influx and reduction in corn seedlings exposed to nitrate and ammonium nutrition.Plant Physiol,1982,62(9):353-359.
[33]Bonser A M,Lynch J,Snapp S.Effect of phosphorus deficiency on growth angle of basal roots in Phaseolus vulgaris.New Phytol,1996,132(2):281-288.
[34]Ingestad T,Agren G I.The Influence of plant nutrition on biomass allocation.Ecol Appl A Publ Ecol Soc America,1991,1(2):168.
[35]Serna M D,Borras R,Legaz F,Primo-Millo E.The influence of nitrogen concentration and ammonium/nitrate ratio on N-uptake,mineral composition and yield of citrus.Plant&Soil,1992,147(1):13-23.
[36]Muchhal U S,Pardo J M,Raghothama K G.Phosphate transporters from the higher plant Arabidopsis thaliana.PNAS,1996,93(19):10519.
[2]Yan X,Liao H,Beebe S E,Blair M W,Lynch J P.QTLmapping of root hair and acid exudation traits and their relationship to phosphorus uptake in common bean.Plant&Soil,2004,265(1/2):17-29.
[3]Bates T R,Lynch J P.Stimulation of root hair elongation in Arabidopsis thaliana by low phosphorus availability.Plant Cell&Environ,1996,19(5):529-538.
[4]Jungk A.Root hairs and the acquisition of plant nutrients from soil.J Plant Nutr Soil Sci,2015,164(2):121-129
[5]Bonser A M,Lynch J,Snapp S.Effect of phosphorus deficiency on growth angle of basal roots in phaseolus Vulgaris.New Phytol,1996,132(2):281-288.
[6]Liao H,Rubio G,Yan X,Cao A Q,Brown K M,Lynch J P.Effect of phosphorus availability on basal root shallowness in common bean.Plant&Soil,2001,232(1/2):69-79
[7]沈宏,严小龙.根系分泌作用及其诱导机制.生态环境学报,2001,10(4):339-342.Sheng H,Yan X L.Root exudation and their induced mechanism.Ecol Environ Sci,2001,10(4):339-342(in Chinese with English abstract)
[8]张平.氮形态对低磷胁迫下苗期玉米磷素吸收的影响及其机理.南京:南京农业大学,2012.Zhang P.Mechanisms and effects of nitrogen forms on phosphorus absorption of maize(Zea mays L.)under phosphorus stress at seedling stage.Nanjing:Nanjing Agricultural University,2012.(in Chinese with English abstract)
[9]邹春琴,杨志福.氮素形态对春小麦根际pH与磷素营养状况的影响.土壤通报,1994(4):175-177.Zhou C Q,Yang Z F.Effects of different form nitrogen on pH and phosphorus content of spring wheat rhizosphere.Chin J Soil Sci,1994(4):175-177.(in Chinese with English abstract)
[10]李朝霞.磷匮乏影响玉米根系发育机制的研究.济南:山东大学,2011.Li C X.Study on the mechanism of root developmental changes to phosphate deficiency in maize.Jinan:Shandong University,2011.(in Chinese with English abstract)
[11]Chen X P,Cui Z L,Vitousek P M,Cassman K G,Matson P A,Bai J S.Integrated soil-crop system management for food security.PNAS,2011,108(16):6399-6404.
[12]严小龙,廖红,戈振扬.植物根构型特性与磷吸收效率.植物学报,2000,17(6):511-519.Yan X L,Liao H,Ge Z Y.root architectural characteristics and phosphorus acquisition efficiency in plants.Chin Bull Bot,2000,17(6):511-519.(in Chinese)
[13]赵静,付家兵,廖红,罗锡文.大豆磷效率应用核心种质的根构型性状评价.科学通报,2004,49(13):1249-1257.Zhao J,Fu J B,Liao H,Luo X W.Properties evaluation of phosphorus efficiency on roots structure of soybean.Chin Sci Bull,2004,49(13):1249-1257.(in Chinese)
[14]Forde B,Lorenzo H.The nutritional control of root development.Interactions in the root environment:An Integrated Approach.2002:51-68.
[15]刘灵,廖红,王秀荣,严小龙.不同根构型大豆对低磷的适应性变化及其与磷效率的关系.中国农业科学,2008,41(4):1089-1099.Liu L,Liao H,Wang X R,Yan X L.Adaptive changes of soybean genotypes with different root architectures to low phosphorus availability as related to phosphorus efficiency.Sci Agric Sin,2008,41(4):1089-1099.(in Chinese with English abstract)
[16]李佳.水稻根系应答根际水分与磷素环境变化的研究.扬州:扬州大学,2015.Li J.Study on rhizosphere moisture and phosphorus environment change of rice root response.Yangzhou:Yangzhou University,2015.(in Chinese with English abstract)
[17]陈晨,龚海青,张敬智,郜红建.水稻根系形态与氮素吸收累积的相关性分析.植物营养与肥料学报,2017,23(2):333-341.Chen C,Gong H Q,Zhang J Z,Gao H J.Correlation between root morphology and nitrogen uptake of rice.J Plant Nutr Fert,2017,23(2):333-341.(in Chinese with English abstract)
[18]Lynch J.Root Architecture and Plant Productivity.Plant Physiol,1995,109(1):7-13.
[19]李宝珍,王松伟,冯慧敏,徐国华.氮素供应形态对水稻根系形态和磷吸收的影响.中国水稻科学,2008,22(6):665-668.Li B Z,Wang S W,Feng H M,Xu G H.Effects of nitrogen forms on root morphology and phosphate uptake in rice.Chin J Rice Sci,2008,22(6):665-668(in Chinese with English abstract)
[20]Ladha J K,Kirk G J,Bennett J,Peng S,Reddy C K,Reddy P M,Singh U.Opportunities for increased nitrogen-use efficiency from improved lowland rice germplasm.Field Crops Res,1998,56(1/2):41-71.
[21]廖红,戈振扬,戈振场,严小龙.水磷耦合胁迫下植物磷吸收的理想根构型:模拟与应用.科学通报,2001,46(8):641-646.Liao H,Ge Z Y,Yan X L,Yan X L.Optimized root architecture for phosphorus acquisition under water and phosphorus stress:Simulation and application.Chin Sci Bull,2001,46(8):641-646.(in Chinese with English abstract)
[22]Muchhal U S,Pardo J M,Raghothama K G.Phosphate transporters from the higher plant Arabidopsis thaliana.PNAS,1996,93(19):10519.
[23]李宝珍,辛伟杰,徐国华.氮饥饿水稻利用不同形态氮素的差异及其生理机制.土壤学报,2007,44(2):273-279.Li B Z,Xin W J,Xu G H.Physiological mechanisms in uptake and use of different forms of nitrogen by nitrogen starved rice crop.Acta Pedol Sin,2007,44(2):273-279.(in Chinese with English abstract)
[24]范志强.氮磷营养及氮形态对水曲柳幼苗生长和生理的影响机制.沈阳:东北林业大学,2004.Fan Z Q.Effects of nitrogen,phosphorus and nitrogen forms on growth and physiological traits of fraxinus mandshurica seedlings.Shenyang:Northeast Forestry University,2004(in Chinese with English abstract)
[25]张彦东,范志强,王庆成,王政权.不同形态N素对水曲柳幼苗生长的影响.应用生态学报,2000,11(5):665-667.Zhou Y D,Fan Z Q,Wang Q C,Wang Z Q.Effects of different nitrogen forms on growth of Fraxinus mandshurica seedlings.Chin J Appl Ecol,2000,11(5):665-667.(in Chinese with English abstract)
[26]Miller M H.Effects of nitrogen on phosphorus absorption by plants.Proc Plant Root Environ,1974.
[27]Thien S J,Mcfee W W.Effect of Nitrogen on phosphorus transport systems in Zea Mays L.Soil Sci Soc Amer J,1972,36(4):617-620.
[28]Smith F W,Jackson W A.Nitrogen enhancement of phosphate transport in roots of Zea mays L.:I.Effects of ammonium and nitrate pretreatment.Plant Physiol,1987,84(4):1314-1318.
[29]Hirel B,Limami A M.Prospects for improving nitrogen use efficiency:Insights given by 15N-labelling experiments.Phytochem Rev,2002,2(1/2):133-144.
[30]Radin J W.Differential regulation of nitrate reductase induction in roots and shoots of cotton plants.Plant Physiol,1975,55(2):178-182.
[31]Zhang H,Xue YG,Wang ZQ,Yang J,Zhang J.Morphological and physiological traits of roots and their relationships with shoot growth in“super”rice.Field Crops Res,2009,113(1):31-40.
[32]Mackown C T,Jackson W A,Volk R J.Restricted nitrate influx and reduction in corn seedlings exposed to nitrate and ammonium nutrition.Plant Physiol,1982,62(9):353-359.
[33]Bonser A M,Lynch J,Snapp S.Effect of phosphorus deficiency on growth angle of basal roots in Phaseolus vulgaris.New Phytol,1996,132(2):281-288.
[34]Ingestad T,Agren G I.The Influence of plant nutrition on biomass allocation.Ecol Appl A Publ Ecol Soc America,1991,1(2):168.
[35]Serna M D,Borras R,Legaz F,Primo-Millo E.The influence of nitrogen concentration and ammonium/nitrate ratio on N-uptake,mineral composition and yield of citrus.Plant&Soil,1992,147(1):13-23.
[36]Muchhal U S,Pardo J M,Raghothama K G.Phosphate transporters from the higher plant Arabidopsis thaliana.PNAS,1996,93(19):10519.
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