玉米根系吸水调控机制
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摘要
根系吸水对于作物生长发育及产量有着重要的影响，深入了解根系吸水机理，分析不同调控措施对根系吸水的影响，对于实现节水、高效、高产的现代农业具有重要的科学价值和现实意义。本论文以玉米为研究对象，采用水培试验和盆栽试验相结合的方法，就不同玉米品种、水分条件和氮素形态调控措施下，作物生长、生理特性、根系吸水能力、水分利用效率等问题进行了研究，以期为作物的高效用水机理提供科学依据。研究主要取得以下结果：
（1）水分条件明显影响了玉米根系形态特征，不同玉米品种间存在显著差异，与天四和478相比，户四具有较大的根系生长。与正常供水相比，干旱胁迫下三个玉米品种根系总表面积和总根长增加，根系干重降低。干旱胁迫显著增加玉米初生根长、次生根长、种子根主根长、种子根总根长、一级侧根密度和平均长度。两种水分条件下，三个玉米品种间的根系生长总体表现为户四优于478和天四的趋势。与其它两个品种相比，户四具有较高的根系干重、较大的总根长和总表面积。而户四较大的总根长主要是由于初生根特别是种子根长较大，一级侧根密度和长度也较大，而次生根长的品种间差异不显著。总体而言，户四的根系生长体现出在适应水分环境方面的优势。
（2）水分条件明显影响了玉米根系吸水能力，不同品种在细胞、单根和整根水平的根系吸水能力存在显著差异，表现为户四根系吸水能力优于天四和478。细胞水平下，水分胁迫显著降低了三个玉米品种的根皮层细胞膨压，表现为户四>天四>478且品种间差异显著，但正常供水条件下品种间差异不显著。用HgCl2和2-巯基乙醇先后处理根皮层细胞，其水导先降低后升高，但并未恢复到初始水平，表明HgCl2抑制了水通道蛋白活性，2-巯基乙醇则使水通道蛋白活性部分恢复。两种水分条件下各处理阶段根皮层细胞水导均表现为户四>天四>478且品种间差异显著，水分胁迫显著降低了根皮层细胞水导。单根水平下，两种水分条件下的玉米静水压下单根水导均为户四>天四>478，品种间差异显著，且水分胁迫显著降低了其径向导度和轴向导度。不同品种在渗透压下也有相同趋势。单根解剖结构与单根水导具有显著的相关性，单根径向导度与皮层厚度占根直径的比例呈负相关关系（R=–0.77, P <0.01）；单根轴向导度则主要受木质部导管的影响，与中央木质部导管直径（R=0.75, P <0.01）和木质部导管总横截面积（R=0.93, P <0.01）呈正相关关系。整根水平下，两种水分条件下三个品种的整根水导呈户四>天四>478的趋势，品种间差异显著，且水分胁迫显著降低了玉米整根水导。
（3）土壤水分条件和氮素形态显著影响了玉米地上部和地下部生长。与正常供水相比，干旱胁迫对玉米地上部和地下部生长均有抑制作用，增加植株根冠比，促进光合作用产物在根部积累。施用氮肥能显著改善玉米生长，且随着施氮量增加效果更显著。相同供氮水平下，硝态氮和铵态氮混施显著提高玉米地上部和地下部生物量，显著降低根冠比，更有利于植株生长，特别是根系的生长发育。水分条件和氮素形态主要通过对玉米净光合速率和叶绿素等因子的调控，从而影响作物光合作用，调控作物生长。
（4）土壤水分条件和氮素形态显著影响了玉米地上部蒸腾耗水和根系吸水能力，从而调控植株体内水分平衡。水分胁迫条件下玉米地上部蒸腾耗水和根系吸水能力较正常供水显著降低。同一供水条件下，施氮能显著提高玉米的地上部蒸腾速率和根系吸水能力，且高氮处理优于低氮处理。硝态氮和铵态氮混施较相同施用量单施处理能更有效增强地上部蒸腾作用，提高生物量水分利用效率；而单施硝态氮可增强细胞和整根水平的根系吸水能力，更有利于使植株保持较好的水分状况。
Water uptake by roots plays an important role in the growth and development aswell as the yield of crops. In-depth understanding of the mechanism water uptake byroots and analysis the impact of different control measures on water uptake by rootare of scientific values and practical significance to water-saving, high-efficiency,high-yielding modern agriculture. To investigate the mechanism of water uptake bycrop roots, this research project chose maize as the plant material and employed acombination method of water culture experiment and pot experiment. The key cropindices, e.g., crop morphogenesis, physiological characteristics, water uptake abilityby roots, and water use efficiency were determined under different control measuresin terms of corn genotype (i.e., Hu4, Tian4, and478), water condition (i.e., wellwatered and water deficit conditions), and nitrogen (N) form of the fertilizer(nitrate-N and ammonium-N). The results should contribute to understanding of themechanism of efficient water use by crops. The main findings of this study were asfollows:
(1) Water supply conditions substantially influenced the morphologicalcharacteristics of maize roots. There were significant variations among differentmaize genotypes. Hu4produced a larger root system than Tian4and478.Compared with well watered, water deficit increased the total root surface area andtotal root length while decreasing the root dry weight of different maize genotypes. Inaddition, water deficit significantly increased the length of primary and secondaryroots, the length and main root length of seminal roots, and the linear frequency andlength of first-order lateral roots. Under different water regimes, the root growth ofHu4was better than that of478and Tian4. As compared to478and Tian4, Hu4 had greater root dry weight, total root length, and total surface area. In particular, Hu4was characterized by large length of primary roots as well as large linear frequencyand length of first-order lateral roots, whereas the length of secondary roots showedno significant variations among different maize genotypes. Overall, the root growth ofHu4showed advantages in adapting to the water conditions.
(2) Water supply conditions substantially influenced water uptake by maizeroots. There were significant variations among different maize genotypes atcellular, single root and whole root system levels, with the water uptake ability ofHu4was higher than that of Tian4and478. At cellular level, the turgor of corticalroot cells of the three maize genotypes decreased under water deficit, and the turgorranking was in the order Hu4> Tian4>478. But, the turgor of the three maizegenotypes was with no genotypic differences under the well watered condition. Aftertreated by HgCl2and2-mercaptoethanol under two water levels，cell hydraulicconductivity (Lpc) of the three maize genotypes were firstly decreased，then increased,and the final values were even lower than the original values. The results indicatedHgCl2inhibited the activity of aquaporins, and2-mercaptoethanol parted reversed theactivity of aquaporins. The Lpcof the three genotypes substantially decreased in thewater deficit treatment, and the Lpcranking was in the order Hu4> Tian4>478inthe two water treatments. At single root level, the hydrostatic hydraulic conductivity(Lpsr) of single roots varied among genotypes under the two water levels, with thehighest in Hu4and the lowest in478. Radial hydraulic conductivity (radial Lpsr) andaxial hydraulic conductance (Lax) of the three genotypes varied similarly as Lpsr, andwater stress decreased radial Lpsrand Lax. The osmotic Lpsrin the Hu4was thehighest (Hu4> Tian4>478) across the water conditions. The variations in hydraulicparameters were related to root anatomy. Radial Lpsrwas negatively correlated withthe ratio of cortex width to root diameter (R=–0.77, P <0.01), whereas Laxwaspositively correlated with the diameter of the central xylem vessel (R=0.75, P <0.01)and the cross-sectional area of xylem vessels (R=0.93, P <0.01). At whole rootsystem level, hydraulic conductivity (Lpwr) of the whole root system followed thesame trend under the two water conditions, with the highest values in the Hu4.Moreover, the Lpwrof each genotype substantially decreased under water deficit.
(3) Soil moisture conditions and N forms significantly influenced the growthof the shoot and root system of maize. Compared with well watered condition,water deficit had an inhibitory effect on the growth of the shoot and root system ofmaize, which increased the root-to-shoot ratio and promoted the accumulation ofproducts of photosynthesis in the roots. Application of N fertilizer significantlyimproved plant growth in a dose-dependent manner. When applied at the same level,the mixture of nitrate-N and ammonium-N significantly increased plant biomass ofthe shoot and root system while significantly reducing the root-to-shoot ratio. Thus,this application of mixed N fertilizers was more conducive to plant growth, especiallythe root growth and development. Soil moisture conditions and N forms mainlythrough regulation of the net photosynthetic rate and the chlorophyll content of maizeleaves, further influencing the photosynthesis process and regulating growth of thecrop.
(4) Soil moisture conditions and N forms significantly influenced thetranspiration and root water uptake capability of the above-ground part ofmaize, thereby regulating water balance in plant body. Under water deficit, thetranspiration and root water uptake capability of the above-ground part of maize weresignificantly lower than those under well watered condition. Under the same waterconditions, application of N fertilizers significantly improved transpiration rate of theabove-ground part and water uptake ability of maize roots, and the improvementeffect was stronger in higher-N treatments than in low-N treatments. Application ofnitrate-N mixed with ammonium-N was more effective than single use of the Nfertilizer in enhancing th e transpiration of the above-ground part of maize andimproving the water use efficiency of the biomass. On the other side, application ofnitrate-N alone enhanced the water uptake ability by roots at cellular and whole rootsystem levels, thus facilitating the plants to maintain good water conditions.
（1）水分条件明显影响了玉米根系形态特征，不同玉米品种间存在显著差异，与天四和478相比，户四具有较大的根系生长。与正常供水相比，干旱胁迫下三个玉米品种根系总表面积和总根长增加，根系干重降低。干旱胁迫显著增加玉米初生根长、次生根长、种子根主根长、种子根总根长、一级侧根密度和平均长度。两种水分条件下，三个玉米品种间的根系生长总体表现为户四优于478和天四的趋势。与其它两个品种相比，户四具有较高的根系干重、较大的总根长和总表面积。而户四较大的总根长主要是由于初生根特别是种子根长较大，一级侧根密度和长度也较大，而次生根长的品种间差异不显著。总体而言，户四的根系生长体现出在适应水分环境方面的优势。
（2）水分条件明显影响了玉米根系吸水能力，不同品种在细胞、单根和整根水平的根系吸水能力存在显著差异，表现为户四根系吸水能力优于天四和478。细胞水平下，水分胁迫显著降低了三个玉米品种的根皮层细胞膨压，表现为户四>天四>478且品种间差异显著，但正常供水条件下品种间差异不显著。用HgCl2和2-巯基乙醇先后处理根皮层细胞，其水导先降低后升高，但并未恢复到初始水平，表明HgCl2抑制了水通道蛋白活性，2-巯基乙醇则使水通道蛋白活性部分恢复。两种水分条件下各处理阶段根皮层细胞水导均表现为户四>天四>478且品种间差异显著，水分胁迫显著降低了根皮层细胞水导。单根水平下，两种水分条件下的玉米静水压下单根水导均为户四>天四>478，品种间差异显著，且水分胁迫显著降低了其径向导度和轴向导度。不同品种在渗透压下也有相同趋势。单根解剖结构与单根水导具有显著的相关性，单根径向导度与皮层厚度占根直径的比例呈负相关关系（R=–0.77, P <0.01）；单根轴向导度则主要受木质部导管的影响，与中央木质部导管直径（R=0.75, P <0.01）和木质部导管总横截面积（R=0.93, P <0.01）呈正相关关系。整根水平下，两种水分条件下三个品种的整根水导呈户四>天四>478的趋势，品种间差异显著，且水分胁迫显著降低了玉米整根水导。
（3）土壤水分条件和氮素形态显著影响了玉米地上部和地下部生长。与正常供水相比，干旱胁迫对玉米地上部和地下部生长均有抑制作用，增加植株根冠比，促进光合作用产物在根部积累。施用氮肥能显著改善玉米生长，且随着施氮量增加效果更显著。相同供氮水平下，硝态氮和铵态氮混施显著提高玉米地上部和地下部生物量，显著降低根冠比，更有利于植株生长，特别是根系的生长发育。水分条件和氮素形态主要通过对玉米净光合速率和叶绿素等因子的调控，从而影响作物光合作用，调控作物生长。
（4）土壤水分条件和氮素形态显著影响了玉米地上部蒸腾耗水和根系吸水能力，从而调控植株体内水分平衡。水分胁迫条件下玉米地上部蒸腾耗水和根系吸水能力较正常供水显著降低。同一供水条件下，施氮能显著提高玉米的地上部蒸腾速率和根系吸水能力，且高氮处理优于低氮处理。硝态氮和铵态氮混施较相同施用量单施处理能更有效增强地上部蒸腾作用，提高生物量水分利用效率；而单施硝态氮可增强细胞和整根水平的根系吸水能力，更有利于使植株保持较好的水分状况。
Water uptake by roots plays an important role in the growth and development aswell as the yield of crops. In-depth understanding of the mechanism water uptake byroots and analysis the impact of different control measures on water uptake by rootare of scientific values and practical significance to water-saving, high-efficiency,high-yielding modern agriculture. To investigate the mechanism of water uptake bycrop roots, this research project chose maize as the plant material and employed acombination method of water culture experiment and pot experiment. The key cropindices, e.g., crop morphogenesis, physiological characteristics, water uptake abilityby roots, and water use efficiency were determined under different control measuresin terms of corn genotype (i.e., Hu4, Tian4, and478), water condition (i.e., wellwatered and water deficit conditions), and nitrogen (N) form of the fertilizer(nitrate-N and ammonium-N). The results should contribute to understanding of themechanism of efficient water use by crops. The main findings of this study were asfollows:
(1) Water supply conditions substantially influenced the morphologicalcharacteristics of maize roots. There were significant variations among differentmaize genotypes. Hu4produced a larger root system than Tian4and478.Compared with well watered, water deficit increased the total root surface area andtotal root length while decreasing the root dry weight of different maize genotypes. Inaddition, water deficit significantly increased the length of primary and secondaryroots, the length and main root length of seminal roots, and the linear frequency andlength of first-order lateral roots. Under different water regimes, the root growth ofHu4was better than that of478and Tian4. As compared to478and Tian4, Hu4 had greater root dry weight, total root length, and total surface area. In particular, Hu4was characterized by large length of primary roots as well as large linear frequencyand length of first-order lateral roots, whereas the length of secondary roots showedno significant variations among different maize genotypes. Overall, the root growth ofHu4showed advantages in adapting to the water conditions.
(2) Water supply conditions substantially influenced water uptake by maizeroots. There were significant variations among different maize genotypes atcellular, single root and whole root system levels, with the water uptake ability ofHu4was higher than that of Tian4and478. At cellular level, the turgor of corticalroot cells of the three maize genotypes decreased under water deficit, and the turgorranking was in the order Hu4> Tian4>478. But, the turgor of the three maizegenotypes was with no genotypic differences under the well watered condition. Aftertreated by HgCl2and2-mercaptoethanol under two water levels，cell hydraulicconductivity (Lpc) of the three maize genotypes were firstly decreased，then increased,and the final values were even lower than the original values. The results indicatedHgCl2inhibited the activity of aquaporins, and2-mercaptoethanol parted reversed theactivity of aquaporins. The Lpcof the three genotypes substantially decreased in thewater deficit treatment, and the Lpcranking was in the order Hu4> Tian4>478inthe two water treatments. At single root level, the hydrostatic hydraulic conductivity(Lpsr) of single roots varied among genotypes under the two water levels, with thehighest in Hu4and the lowest in478. Radial hydraulic conductivity (radial Lpsr) andaxial hydraulic conductance (Lax) of the three genotypes varied similarly as Lpsr, andwater stress decreased radial Lpsrand Lax. The osmotic Lpsrin the Hu4was thehighest (Hu4> Tian4>478) across the water conditions. The variations in hydraulicparameters were related to root anatomy. Radial Lpsrwas negatively correlated withthe ratio of cortex width to root diameter (R=–0.77, P <0.01), whereas Laxwaspositively correlated with the diameter of the central xylem vessel (R=0.75, P <0.01)and the cross-sectional area of xylem vessels (R=0.93, P <0.01). At whole rootsystem level, hydraulic conductivity (Lpwr) of the whole root system followed thesame trend under the two water conditions, with the highest values in the Hu4.Moreover, the Lpwrof each genotype substantially decreased under water deficit.
(3) Soil moisture conditions and N forms significantly influenced the growthof the shoot and root system of maize. Compared with well watered condition,water deficit had an inhibitory effect on the growth of the shoot and root system ofmaize, which increased the root-to-shoot ratio and promoted the accumulation ofproducts of photosynthesis in the roots. Application of N fertilizer significantlyimproved plant growth in a dose-dependent manner. When applied at the same level,the mixture of nitrate-N and ammonium-N significantly increased plant biomass ofthe shoot and root system while significantly reducing the root-to-shoot ratio. Thus,this application of mixed N fertilizers was more conducive to plant growth, especiallythe root growth and development. Soil moisture conditions and N forms mainlythrough regulation of the net photosynthetic rate and the chlorophyll content of maizeleaves, further influencing the photosynthesis process and regulating growth of thecrop.
(4) Soil moisture conditions and N forms significantly influenced thetranspiration and root water uptake capability of the above-ground part ofmaize, thereby regulating water balance in plant body. Under water deficit, thetranspiration and root water uptake capability of the above-ground part of maize weresignificantly lower than those under well watered condition. Under the same waterconditions, application of N fertilizers significantly improved transpiration rate of theabove-ground part and water uptake ability of maize roots, and the improvementeffect was stronger in higher-N treatments than in low-N treatments. Application ofnitrate-N mixed with ammonium-N was more effective than single use of the Nfertilizer in enhancing th e transpiration of the above-ground part of maize andimproving the water use efficiency of the biomass. On the other side, application ofnitrate-N alone enhanced the water uptake ability by roots at cellular and whole rootsystem levels, thus facilitating the plants to maintain good water conditions.
引文
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