Effects of nitrogen application on physiological characteristics and nitrogen uptake and utilization of watermelon under chloride stress
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摘要: 由于长期使用地下含氯微咸水补灌, 氯盐胁迫已经成为限制宁夏西瓜产量和品质的主要因素之一, 而施用氮肥可在一定程度上缓解盐胁迫引起的生长抑制作用, 因此, 探索氮素对西瓜氯盐胁迫的调控机制, 对氯盐胁迫下合理施用氮肥和西瓜氯毒害调控具有重要意义。本研究以‘金城5号’西瓜品种为供试作物, 采用土培试验, 探索氯盐胁迫[160 mg (Cl−1)∙kg−1(烘干土)]下不同施氮水平[0 g∙kg−1(烘干土)、0.10 g∙kg−1(烘干土)、0.15 g∙kg−1(烘干土)、0.20 g∙kg−1(烘干土)、0.25 g∙kg−1(烘干土)]对西瓜幼苗阴阳离子平衡、有机渗透调节物质、抗氧化酶活性、氧化损伤和氮素吸收利用的影响, 以期为揭示氮素对作物氯盐胁迫的调控机理提供理论依据。结果表明, 施用氮肥使西瓜根、茎、叶中的Cl−和Na+均显著减少, 而NO3−和K+均显著增加, 因此, 整株Cl−/NO3−值和Na+/K+值分别比不施氮降低46.0%~69.5%和31.0%~54.3%; 叶片中可溶性糖和脯氨酸含量、超氧化物歧化酶和过氧化氢酶活性均在0.15 g∙kg−1氮水平时达最大值, 分别比不施氮提高75.6%、70.1%、55.8%和54.8%, 而丙二醛含量则比不施氮降低59.3%; 同时, 施氮0.15 g∙kg−1时, 西瓜的氮累积量增加157.7%, 硝酸还原酶活性提高62.4%, 氮吸收效率和氮素利用效率分别达26.25%和97.10%, 西瓜植株鲜重和干物质累积量亦显著提高96.9%和29.0%。对施氮量与西瓜各生长生理指标的聚类分析和相关分析表明, 施氮处理对氯盐胁迫的缓解效果表现为0.15 g∙kg−1>0.20 g∙kg−1>0.10 g∙kg−1>0.25 g∙kg−1, 生物量和干物质累积与氮吸收利用效率及氮累积呈显著正相关, 而氮累积量与抗氧化酶活性、渗透调节物质含量间均具有显著的正相关性, 与Na+/K+值、Cl−/ NO3−值和丙二醛含量呈负相关。综合各指标与施氮量之间的曲线拟合结果, 氯盐浓度为160 mg (Cl−1)∙kg−1(烘干土)时西瓜生长和生理活性适宜施氮量为0.14~0.18 g∙kg−1。可见, 氯盐胁迫下适量施氮可通过调节Na+/K+值和Cl−/NO3−值来维持植株体内离子稳态, 并提高渗透调节物质含量和抗氧化酶活性, 从而降低细胞膜氧化损伤, 增强西瓜植株的生理抗性, 达到对氯盐胁迫的调控作用。Abstract: Chloride stress is one of the main factors limiting the yield and quality of watermelon in Ningxia due to long-term irrigation with underground chlorinated brackish water. Nitrogen could alleviate the chlorine toxicity of crops. Therefore, it is crucial to explore the regulatory mechanism of nitrogen on the chloride stress by applying nitrogen fertilizer rationally and controlling chlorosis of watermelon under chloride stress. In this study, a soil culture experiment was conducted to determine the effects of different nitrogen application rates [0, 0.10, 0.15, 0.20, 0.25 g∙kg−1 (oven-dry soil)] on anion-cation balance, organic osmotic regulators, antioxidant enzyme activity, oxidative damage, and nitrogen uptake and utilization in watermelon seedlings under chloride stress of 160 mg(Cl−1)∙kg−1 (oven-dry soil). The test crop was the ‘Jincheng No. 5’ variety of watermelon. The results showed that nitrogen application considerably reduced Cl− and Na+ contents in the roots, stems, and leaves of watermelon while significantly increased NO3− and K+ contents at P<0.05; thus the ratios of Cl−/NO3− and Na+/K+ of the whole plant decreased by 46.0%−69.5% and 31.0%−54.3% compared with that of 0 g∙kg−1 nitrogen rate, respectively. Moreover, the contents of soluble sugar and proline, the activities of superoxide dismutase (SOD) and catalase (CAT) in leaves all reached the maximum levels at 0.15 g∙kg−1 N, which statistically increased by 75.6%, 70.1%, 55.8%, and 54.8% at P<0.05 compared with those at 0 g∙kg−1 N, respectively; while the content of malondialdehyde (MDA) significantly decreased by 59.3%. Moreover, when nitrogen was applied at 0.15 g∙kg−1, the nitrogen accumulation of watermelon increased by 157.7%, activity of nitrate reductase (NR) increased by 62.4%, and nitrogen uptake efficiency and nitrogen use efficiency were 26.25% and 97.10%, respectively. Thus, the fresh and dry weights of the plant increased by 96.9% and 29.0% at P<0.05, respectively. Cluster and correlation analyses of nitrogen application rate and physiological growth indexes of watermelon showed that the mitigation effect of nitrogen application on watermelon chloride stress was 0.15 g∙kg−1 > 0.20 g∙kg−1 > 0.10 g∙kg−1 > 0.25 g∙kg−1. There was significant positive correlation between biomass and dry matter accumulation with nitrogen uptake, utilization efficiency, and nitrogen accumulation; while there were also significant positive correlations between nitrogen accumulation and antioxidant enzyme activity and osmo-modulator content, and negative correlations with Na+/K+ ratio, Cl−/NO3− ratio, and MDA content. Based on the curve fitting results of each index, the nitrogen application rate of 0.14−0.18 g∙kg−1 was available for the growth and physiological activity of watermelon when the chloride concentration was 160 mg(Cl−1)∙kg−1(oven-dry soil). This indicates that appropriate nitrogen application under chloride salt stress can maintain ion homeostasis in plants by adjusting the Na+/K+ and Cl−/NO3− ratios, as well as improve the content of osmoregulatory substances and antioxidant enzyme activities, thereby reducing cell membrane oxidative damage, enhancing the physiological resistance of watermelon plants, and achieving a regulatory effect on chloride stress.
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图 1 氯胁迫下施氮水平对西瓜幼苗生物量的影响
Figure 1. Effects of N rate on biomass of watermelon under chlorine stress
图 2 氯胁迫下施氮水平对西瓜幼苗渗透调节物质含量的影响
Figure 2. Effect of N rate on organic osmoregulation substances contents of watermelon seedlings under chlorine stress
图 3 氯胁迫下施氮水平对西瓜幼苗抗氧化酶(SOD: 超氧化物歧化酶; CAT: 过氧化氢酶)活性的影响
Figure 3. Effect of N rate on antioxidant enzymes (SOD: superoxide dismutase; CAT: catalase) activities of watermelon seedlings under chlorine stress
图 4 氯胁迫下施氮水平对西瓜幼苗丙二醛(MDA)含量的影响
Figure 4. Effect of N rate on malondialdehyde (MDA) content of watermelon seedlings under chlorine stress
图 5 氯胁迫下不同氮水平处理西瓜幼苗相关指标聚类分析热图和相关分析图
FW: 生物量; DM: 干物质; Pro: 脯氨酸含量; ss: 可溶性糖含量; SOD: 超氧化物歧化酶活性; CAT: 过氧化氢酶活性; MDA: 丙二醛含量; N Accum: 氮累积; N Abs: 氮吸收效率; NUE: 氮利用率; NR: 硝酸还原酶活性。Na/K(L): leaf Na+/K+ ratio; Na/K(S): stem Na+/K+ ratio; Na/K(R): root Na+/K+ ratio; Cl/N(L): leaf Cl−/NO3− ratio; Cl/N(S): stem Cl−/NO3− ratio; Cl/N(R): root Cl−/NO3− ratio; FW: frest weight; DM: dry matter; Pro: proline content; ss: soluble sugar content; SOD: superoxide dismutase activity; CAT: catalase activity; MDA: malondialdehyde content; N Accum: N accumulation; N Abs: N absorption efficiency; NUE: N utilization ratio; NR: nitrate reductase activity.
Figure 5. Heat map and correlation analysis diagram among relative indexes of watermelon seedlings of different N rates under chlorine stress
表 1 供试土壤化学性质
Table 1. Soil basic physical and chemical properties
指标
Index
pH有机质
Organic matter
(g∙kg−1)全氮
Total nitrogen
(g∙kg−1)全磷
Total phosphorus
(g∙kg−1)矿质态氮
Mineral nitrogen
(mg∙kg−1)速效磷
Available phosphorus
(mg∙kg−1)速效钾
Available potassium
(mg∙kg−1)氯离子
Chloridion
(g∙kg−1)数值 Value 8.03 7.14 0.96 0.87 31.56 6.95 114.88 0.06 
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表 2 氯胁迫下不同施氮水平对西瓜幼苗离子含量的影响
Table 2. Ions contents of watermelon seedlings affected by N rate under chlorine stress
项目
Item氮水平 N rate (g∙kg−1) 0 0.10 0.15 0.20 0.25 K+含量
K+ content
(g∙kg−1)叶 Leaf 7.08±0.81c 8.10±0.82bc 9.44±0.94ab 10.44±0.47a 7.43±0.47bc 茎 Stem 6.43±0.47b 8.77±0.94ab 9.10±0.82ab 10.44±1.70a 8.77±1.25ab 根 Root 2.09±0.82b 2.75±0.47ab 3.76±0.72ab 4.43±0.72a 2.42±0.47b 整株 Plant 5.20±0.31c 6.54±0.61bc 7.32±1.14ab 8.43±0.55a 6.32±0.16bc Na+含量
Na+ content
(g∙kg−1)叶 Leaf 0.32±0.03a 0.21±0.03b 0.17±0.02b 0.17±0.02b 0.19±0.02b 茎 Stem 0.44±0.05a 0.40±0.03ab 0.38±0.03b 0.39±0.03ab 0.43±0.01ab 根 Root 0.70±0.03a 0.58±0.03abc 0.46±0.04c 0.52±0.03bc 0.61±0.05ab 整株 Plant 0.49±0.01a 0.40±0.01b 0.33±0.01c 0.36±0.02bc 0.41±0.03b Na+/K+ 叶 Leaf 0.05a 0.03b 0.02b 0.02b 0.03b 茎 Stem 0.07a 0.05ab 0.04b 0.04b 0.05ab 根 Root 0.34a 0.21ab 0.12b 0.12b 0.25ab 整株 Plant 0.09a 0.06bc 0.05c 0.04c 0.06b NO3−含量
NO3− content
(mg∙kg−1)叶 Leaf 230.4±30.9b 239.6±14.4b 287.0±16.2ab 314.5±17.3a 305.4±19.6ab 茎 Stem 273.2±10.0b 355.8±8.7b 415.4±6.2b 754.9±10.6a 377.2±12.5b 根 Root 152.4±26.2b 337.5±42.0a 366.5±14.2a 430.7±42.6a 267.1±26.1ab 整株 Plant 218.7±17.7c 311.0±25.2bc 356.3±6.9b 500.1±69.8a 316.6±4.4b Cl−含量
Cl− content
(mg∙kg−1)叶 Leaf 8878.6±397.6a 6418.6±389.3ab 4921.4±432.5b 6209.1±587.9b 6284.3±438.0ab 茎 Stem 12 866.7±599.4a 11 011.0±635.9ab 9950.2±809.1b 10 071.0±488.2b 11 553.5±231.4ab 根 Root 7882.3±266.8a 5324.2±274.6b 4004.9±234.4b 4355.2±377.3b 5089.2±272.8b 整株 Plant 9875.9±183.7a 7584.6±457.4b 6292.2±853.3b 6878.4±781.6b 7642.3±283.4b Cl−/NO3− 叶 Leaf 38.53±5.95a 26.79±5.61ab 17.15±1.32b 19.74±4.55ab 20.58±3.48ab 茎 Stem 47.09±3.21a 30.95±1.65a 23.95±2.46ab 13.34±3.03b 30.63±1.42a 根 Root 51.71±6.12a 15.78±1.16b 10.93±1.04b 10.11±1.40b 19.05±1.24b 整株 Plant 45.16±3.93a 24.39±3.51b 17.66±2.22bc 13.76±3.65c 24.14±0.71bc 不同小写字母表示施氮水平间在P<0.05水平差异显著。Different lowercase letters mean significant differences among different nitrogen rates at P<0.05 level.
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表 3 氯胁迫下不同施氮水平对西瓜幼苗氮吸收利用的影响
Table 3. Effects of different N rates on N uptake and utilization of watermelon seedlings under chlorine stress
氮水平
N rate (g∙kg−1)氮吸收累积量
N absorption accumulation (g∙plant−1)氮吸收效率
N absorption efficiency
(%)氮利用率
N utilization ratio
(%)硝酸还原酶活性
Nitrate reductase activity
(μg∙g−1)叶
Leaf茎
Stem根
Root整株
Whole plant0 4.23±0.15d 4.87±0.21b 1.27±0.15c 10.38±0.24d 11.00±0.78b — 12.26±1.40c 0.10 8.64±1.51c 6.63±1.50ab 1.76±0.18b 17.03±2.27c 16.04±1.31ab 66.50±17.35ab 16.27±1.82bc 0.15 15.25±1.16a 9.21±1.04a 2.28±0.06a 26.75±0.88a 26.25±2.54a 97.10±7.86a 19.91±1.43ab 0.20 13.70±1.58ab 8.76±0.56a 2.26±0.19a 24.72±2.24ab 26.71±2.23a 71.73±6.72ab 21.91±1.38a 0.25 11.00±1.50bc 7.01±1.12ab 2.27±0.07a 20.27±2.60bc 24.10±3.94ab 39.56±6.01b 15.69±0.76bc 不同小写字母表示施氮水平间在P<0.05水平差异显著。Different lowercase letters mean significant differences among different nitrogen rates at P<0.05 level.
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