Effects of water and nitrogen regulation on physiological characteristics and yield of cotton
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摘要: 水分和氮素是影响棉花生长发育和产量的主要因子, 为探究水分和氮素对棉花形态、生理特性以及产量的调控效应, 本研究以‘农大棉36号’为材料, 设置干旱胁迫(W1, 相对含水量为45%±5%)和正常供水(W2, 相对含水量为70%±5%)两个水分条件以及不施氮肥(N0)、低氮[N1, 69 mg(N)∙kg−1]、常规施氮[N2, 138 mg(N)∙kg−1] 3个氮素水平, 分析不同水分和氮肥条件下棉花地上部和根系形态、光合性状、抗氧化酶活性、氮代谢酶活性以及产量的变化。结果表明, 与W2处理相比, W1处理显著抑制了棉花生长, 降低了棉花株高、茎粗、叶面积以及总根长、总根表面积、平均根直径(P<0.05), 显著增加了超氧化物歧化酶(SOD)、过氧化物酶(POD)、过氧化氢酶(CAT)活性, 降低了相对叶绿素含量(SPAD)、最大光化学效率(Fv/Fm)和光合能力, 从而造成棉花产量下降(P<0.05); 与N2处理相比, N0和N1处理上述指标均显著降低。干旱胁迫下, 常规施氮处理比不施氮肥和低氮处理促进棉花地上和地下生长, 显著提高主茎叶SPAD、净光合速率和Fv/Fm (P<0.05), 增强抗氧化酶(SOD、POD、CAT)和氮代谢酶(GS和NR)活性(P<0.05), 减轻干旱胁迫对棉花生长造成的抑制, 提高棉花产量。在低氮条件下, 正常供水比干旱胁迫处理亦促进棉花的生长, 增强光合作用和氮代谢酶活性, 减缓了低氮胁迫对其产生的不利影响, 提高产量(P<0.05)。因此, 可通过增施氮肥提高干旱胁迫下的棉花产量, 亦可通过适当增加灌水量提高低氮胁迫下的棉花产量, 本研究为明确氮素胁迫和水分胁迫下合理的水肥管理提供了理论依据。Abstract: Water and nitrogen are the main factors affecting the growth, development, and yield of cotton. In this study, “Nongda Cotton No. 36” was selected to investigate the effects of water and nitrogen on morphology, physiological characteristics, and yield of cotton. Two water conditions were set: drought stress (W1, relative water content was 45%±5%) and normal water supply (W2, relative water content was 70%±5%), and three nitrogen levels: no nitrogen (N0), low nitrogen [N1, 69 mg(N)∙kg−1], and normal nitrogen fertilizers [N2, 138 mg(N)∙kg−1]. Changes in aboveground and root morphology, photosynthetic characteristics, antioxidant enzyme activity, nitrogen metabolism enzyme activity, and cotton yield were analyzed under different water and nitrogen fertilizer conditions. The results showed that compared with the W2 treatments, the W1 treatments significantly inhibited cotton growth and decreased plant height, stem diameter, leaf area, total root length, total root surface area, and average root diameter (P<0.05). The activities of superoxide dismutase (SOD), peroxidase (POD), and catalase (CAT) were significantly enhanced, the relative chlorophyll content (SPAD) and maximum photochemical efficiency (Fv/Fm) were decreased, and the photosynthetic capacity was weakened, resulting in a decrease in cotton yield (P<0.05). Compared with the N2 treatments, the N0 and N1 treatments significantly reduced plant height, stem diameter, leaf area, total root length, total root surface area, and average root diameter of cotton (P<0.05). The activities of SOD, POD, CAT, glutamine synthetase (GS), nitrate reductase (NR), SPAD, and Fv/Fm in cotton were significantly decreased under the N0 and N1 treatments, and the photosynthetic capacity of cotton was weakened. Thus, the cotton yield decreased (P<0.05). Under drought stress, conventional nitrogen application promoted the growth of the aboveground and underground parts of cotton; significantly increased the SPAD, net photosynthetic rate, and Fv/Fm in main-stem leaves (P<0.05); and enhanced the activities of antioxidant enzymes (SOD, POD, and CAT) and nitrogen metabolism enzymes (GS and NR) (P<0.05), which alleviated the damage caused by drought stress and increased the cotton yield. Under low-nitrogen conditions, the normal water supply treatment promoted the growth of cotton; enhanced photosynthesis, nitrogen metabolism enzyme activities, and yield (P<0.05); and alleviated the adverse effects of low-nitrogen stress on cotton. Therefore, cotton yield under drought stress can be increased by increasing nitrogen fertilizer, and cotton yield under low-nitrogen stress can be increased by appropriately increasing irrigation water. The results provide a theoretical basis for clarifying rational water and fertilizer management of cotton under nitrogen and water stresses.
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图 1 水分和氮素对棉花株高(A)、茎粗(B)以及叶面积(C)的影响
W1: 相对含水量为45%±5%; W2: 相对含水量为70%±5%; N0: 不施氮肥; N1: 施氮69 mg(N)·kg−1; N2: 施氮138 mg(N)·kg−1。不同小写字母表示同一时间不同处理间在P<0.05水平差异显著。ns表示未达到显著水平, *代表P<0.05显著水平, **代表达P<0.01极显著水平。W1: relative water content of 45%±5%; W2: relative water content of 70%±5%; N0: no nitrogen fertilizer; N1: nitrogen application rate of 69 mg(N)·kg−1; N2: nitrogen application rate of 138 mg(N)·kg−1. Different lowercase letters above the bars indicate significant differences at P<0.05 level among treatments in the same date. ns means no significant difference. * means signifi-cant differences at P<0.05 level. ** means significant differences at P<0.01 level.
Figure 1. Effects of water and nitrogen regulation on plant height (A), stem diameter (B) and leaf area (C) of cotton
图 2 水分和氮素调控对棉花总根长(A)、总根表面积(B)以及平均根直径(C)的影响
W1: 相对含水量为45%±5%; W2: 相对含水量为70%±5%; N0: 不施氮肥; N1: 施氮69 mg(N)·kg−1; N2: 施氮138 mg(N)·kg−1。不同小写字母表示不同处理间在P<0.05水平差异显著。ns表示未达到显著水平, *代表P<0.05显著水平, **代表P<0.01极显著水平。W1: relative water content of 45%±5%; W2: relative water content of 70%±5%; N0: no nitrogen fertilizer; N1: nitrogen application rate of 69 mg(N)·kg−1; N2: nitrogen application rate of 138 mg(N)·kg−1. Different lowercase letters above the bars indicate significant differences at P<0.05 level among treatments. ns means no significant difference. * means significant differences at P<0.05 level. ** means significant differences at P<0.01 level.
Figure 2. Effects of water and nitrogen regulation on total root length (A), total root surface area (B) and average root diameter (C) of cotton
图 3 水分和氮素调控对棉花谷氨酰胺合成酶活性(A)和硝酸还原酶活性(B)的影响
W1: 相对含水量为45%±5%; W2: 相对含水量为70%±5%; N0: 不施氮肥; N1: 施氮69 mg(N)·kg−1; N2: 施氮138 mg(N)·kg−1。不同小写字母表示同一时间不同处理间在P<0.05水平差异显著。ns表示未达到显著水平, *代表P<0.05显著水平, **代表P<0.01极显著水平。W1: relative water content of 45%±5%; W2: relative water content of 70%±5%; N0: no nitrogen fertilizer; N1: nitrogen application rate of 69 mg(N)·kg−1; N2: nitrogen application rate of 138 mg(N)·kg−1. Different lowercase letters above the bars indicate significant differences at P<0.05 level among treatments in the same date. ns means no significant difference. * means significant differences at P<0.05 level. ** means significant differences at P<0.01 level.
Figure 3. Effects of water and nitrogen regulation on activities of glutamine synthetase (GS, A) and nitrate reductase (NR, B) in cotton
图 4 水分和氮素调控对棉花籽棉产量的影响
W1: 相对含水量为45%±5%; W2: 相对含水量为70%±5%; N0: 不施氮肥; N1: 施氮69 mg(N)·kg−1; N2: 施氮138 mg(N)·kg−1。不同小写字母表示不同处理间在P<0.05水平差异显著。ns表示未达到显著水平, *代表P<0.05显著水平, **代表P<0.01极显著水平。W1: relative water content of 45%±5%; W2: relative water content of 70%±5%; N0: no nitrogen fertilizer; N1: nitrogen application rate of 69 mg(N)·kg−1; N2: nitrogen application rate of 138 mg(N)·kg−1. Different lowercase letters above the bars indicate significant differences at P<0.05 level among treatments. ns means no significant difference. * means significant differences at P<0.05 level. ** means significant differences at P<0.01 level.
Figure 4. Effects of water and nitrogen regulation on cotton seed yield
表 1 水分和氮素调控对棉花相对叶绿素含量(SPAD)的影响
Table 1. Effects of water and nitrogen regulation on relative chlorophyll content (SPAD) of cotton
处理
Treatment处理后天数 Days after treatment (d) 0 15 30 45 60 75 90 W1N0 42.33±0.50a 42.97±0.85e 41.20±0.26d 35.63±0.75e 30.87±0.68e 25.73±1.37e 20.33±1.25d W2N0 43.07±0.95a 46.00±0.46cd 43.67±0.59c 40.70±0.85c 35.87±0.85d 32.47±1.86d 28.43±0.68c W1N1 42.57±0.74a 45.10±0.26d 42.30±1.01cd 38.67±0.38d 34.93±0.32d 31.23±1.23d 26.37±0.87c W2N1 43.00±1.15a 47.53±0.95b 50.77±0.57b 45.60±0.46b 42.40±0.60b 40.53±1.19b 36.97±1.16b W1N2 43.10±0.70a 46.63±0.65bc 49.27±1.04b 44.27±0.95b 40.67±0.50c 37.77±0.99c 34.77±0.81b W2N2 42.90±1.11a 49.33±0.93a 55.27±0.95a 50.53±0.86a 47.43±0.47a 44.67±1.20a 41.90±1.41a 水分 Water (W) ns ** ** ** ** ** ** 施氮 Nitrogen application (N) ns ** ** ** ** ** ** 水分×氮肥 W×N ns ns ** * * ns * W1: 相对含水量为45%±5%; W2: 相对含水量为70%±5%; N0: 不施氮肥; N1: 施氮69 mg(N)·kg−1; N2: 施氮138 mg(N)·kg−1。同列不同小写字母表示同一时间不同处理间在P<0.05水平差异显著。ns表示未达到显著水平, *代表P<0.05显著水平, **代表P<0.01极显著水平。W1: relative water content of 45%±5%; W2: relative water content of 70%±5%; N0: no nitrogen fertilizer; N1: nitrogen application rate of 69 mg(N)·kg−1; N2: nitrogen application rate of 138 mg(N)·kg−1. Different lowercase letters in the same column indicate significant differences at P<0.05 level among treatments in the same date. ns means no significant difference. * means significant differences at P<0.05 level. ** means significant differences at P<0.01 level. 
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表 2 水分和氮素调控对棉花净光合速率的影响
Table 2. Effects of water and nitrogen regulation on net photosynthetic rate of cotton
µmol·m−2·s−1 处理
Treatment处理后天数 Days after treatment (d) 0 15 30 45 60 75 90 W1N0 11.65±0.66a 11.04±0.20d 10.31±0.19d 9.99±0.47c 8.94±0.36d 8.21±0.18d 7.79±0.15c W2N0 11.61±0.70a 11.73±0.21c 11.06±0.14c 10.65±0.37bc 9.88±0.14c 9.54±0.52c 9.19±0.04b W1N1 11.90±0.15a 11.42±0.40cd 10.97±0.26c 10.50±0.45bc 9.73±0.08c 9.43±0.19c 9.08±0.11b W2N1 11.99±0.29a 12.37±0.36b 11.96±0.20b 11.17±0.17b 10.58±0.30b 10.14±0.08b 9.63±0.50b W1N2 12.09±0.40a 11.79±0.33c 11.30±0.33c 10.96±0.40b 10.37±0.37b 10.04±0.07b 9.51±0.39b W2N2 12.13±0.19a 13.51±0.46a 13.10±0.40a 12.51±0.26a 11.87±0.21a 11.20±0.25a 10.65±0.22a 水分 Water (W) ns ** ** ** ** ** ** 施氮 Nitrogen application (N) ns ** ** ** ** ** ** 水分×氮肥 W×N ns * * ns * ns ns W1: 相对含水量为45%±5%; W2: 相对含水量为70%±5%; N0: 不施氮肥; N1: 施氮69 mg(N)·kg−1; N2: 施氮138 mg(N)·kg−1。同列不同小写字母表示同一时间不同处理间在P<0.05水平差异显著。ns表示未达到显著水平, *代表P<0.05显著水平, **代表P<0.01极显著水平。W1: relative water content of 45%±5%; W2: relative water content of 70%±5%; N0: no nitrogen fertilizer; N1: nitrogen application rate of 69 mg(N)·kg−1; N2: nitrogen application rate of 138 mg(N)·kg−1. Different lowercase letters in the same column indicate significant differences at P<0.05 level among treatments in the same date. ns means no significant difference. * means significant differences at P<0.05 level. ** means significant differences at P<0.01 level.
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表 3 水分和氮素调控对棉花PSⅡ反应中心光能转换效率(Fv/Fm)的影响
Table 3. Effects of water and nitrogen regulation on light energy conversion efficiency of PSⅡ reaction center (Fv/Fm) of cotton
处理
Treatment处理后天数 Days after treatment (d) 0 15 30 45 60 75 90 W1N0 0.78±0.01a 0.72±0.003e 0.66±0.003e 0.55±0.003e 0.48±0.007d 0.48±0.007d 0.45±0.013d W2N0 0.79±0.012a 0.75±0.005cd 0.69±0.003c 0.59±0.004cd 0.52±0.011c 0.52±0.011c 0.48±0.004c W1N1 0.79±0.014a 0.73±0.003de 0.69±0.003d 0.58±0.014d 0.52±0.012c 0.52±0.012c 0.48±0.007bc W2N1 0.79±0.014a 0.76±0.003b 0.71±0.003b 0.62±0.013b 0.54±0.001b 0.54±0.001b 0.49±0.003bc W1N2 0.79±0.008a 0.74±0.014bc 0.70±0.002b 0.61±0.009bc 0.53±0.005bc 0.53±0.005bc 0.50±0.007b W2N2 0.79±0.008a 0.80±0.006a 0.75±0.009a 0.67±0.006a 0.64±0.008a 0.64±0.008a 0.61±0.011a 水分 Water (W) ns ** ** ** ** ** ** 施氮 Nitrogen application (N) ns ** ** ** ** ** ** 水分×氮肥 W×N ns * ** * ** ** ** W1: 相对含水量为45%±5%; W2: 相对含水量为70%±5%; N0: 不施氮肥; N1: 施氮69 mg(N)·kg−1; N2: 施氮138 mg(N)·kg−1。同列不同小写字母表示同一时间不同处理间在P<0.05水平差异显著。ns表示未达到显著水平, *代表P<0.05显著水平, **代表P<0.01极显著水平。W1: relative water content of 45%±5%; W2: relative water content of 70%±5%; N0: no nitrogen fertilizer; N1: nitrogen application rate of 69 mg(N)·kg−1; N2: nitrogen application rate of 138 mg(N)·kg−1. Different lowercase letters in the same column indicate significant differences at P<0.05 level among treatments in the same date. ns means no significant difference. * means significant differences at P<0.05 level. ** means significant differences at P<0.01 level.
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表 4 水分和氮素调控对棉花超氧化物歧化酶(SOD)活性的影响
Table 4. Effects of water and nitrogen regulation on superoxide dismutase (SOD) activity of cotton
U·g−1(FW) 处理
Treatment处理后天数 Days after treatment (d) 0 15 30 45 60 75 90 W1N0 503.84±25.56a 704.81±10.82c 867.84±30.86bc 811.62±19.32c 737.84±11.16c 686.54±21.64c 644.38±27.19c W2N0 510.86±36.29a 645.78±26.02d 808.11±48.36d 749.08±12.35d 699.89±21.39d 642.27±12.35d 608.54±14.81c W1N1 523.51±34.79a 756.81±22.80b 916.32±8.52ab 863.62±14.96b 801.78±11.99b 756.11±21.12b 700.59±14.35b W2N1 527.03±35.96a 702.00±17.26c 862.22±22.21c 812.32±12.71c 746.97±12.88c 706.92±27.51c 657.03±27.51bc W1N2 547.41±11.99a 801.08±17.26a 957.08±19.32a 926.16±13.55a 872.76±21.08a 818.65±19.13a 758.22±37.02a W2N2 548.81±36.66a 754.00±14.04b 910.70±15.20abc 881.89±24.43b 829.89±25.33b 763.14±26.91b 706.22±16.86b 水分 Water (W) ns ** ** ** ** ** ** 施氮 Nitrogen application (N) ns ** ** ** ** ** ** 水分×氮肥 W×N ns ns ns ns ns ns ns W1: 相对含水量为45%±5%; W2: 相对含水量为70%±5%; N0: 不施氮肥; N1: 施氮69 mg(N)·kg−1; N2: 施氮138 mg(N)·kg−1。同列不同小写字母表示同一时间不同处理间在P<0.05水平差异显著。ns表示未达到显著水平, *代表P<0.05显著水平, **代表P<0.01极显著水平。W1: relative water content of 45%±5%; W2: relative water content of 70%±5%; N0: no nitrogen fertilizer; N1: nitrogen application rate of 69 mg(N)·kg−1; N2: nitrogen application rate of 138 mg(N)·kg−1. Different lowercase letters in the same column indicate significant differences at P<0.05 level among treatments in the same date. ns means no significant difference. * means significant differences at P<0.05 level. ** means significant differences at P<0.01 level.
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表 5 水分和氮素调控对棉花过氧化物酶(POD)活性的影响
Table 5. Effects of water and nitrogen regulation on peroxidase (POD) activity of cotton
U·mg−1(FW) 处理
Treatment处理后天数 Days after treatment (d) 0 15 30 45 60 75 90 W1N0 297.45±18.73a 407.41±13.26cd 536.46±28.69c 493.06±27.94c 401.04±30.12b 357.64±18.13c 318.29±18.24c W2N0 300.93±20.12a 375.58±19.12d 483.22±17.82d 435.76±35.87d 343.17±22.12c 299.77±16.41d 278.36±22.39d W1N1 309.61±20.12a 446.18±18.13b 573.50±25.42bc 534.72±22.77abc 438.08±25.42b 402.78±19.33b 344.33±22.59bc W2N1 312.50±15.62a 418.40±15.14bc 543.98±33.14bc 503.47±15.91bc 406.25±27.29b 350.69±21.05c 309.61±21.43cd W1N2 314.81±32.26a 484.95±24.82a 626.16±19.12a 568.87±19.12a 506.37±28.07a 451.39±26.95a 386.00±15.66a W2N2 318.29±12.56a 451.39±16.56ab 583.91±15.17b 541.67±19.1ab 444.44±21.12b 408.56±11.82b 359.95±26.12ab 水分 Water (W) ns ** ** ** ** ** ** 施氮 Nitrogen application (N) ns ** ** ** ** ** ** 水分×氮肥 W×N ns ns ns ns ns ns ns W1: 相对含水量为45%±5%; W2: 相对含水量为70%±5%; N0: 不施氮肥; N1: 施氮69 mg(N)·kg−1; N2: 施氮138 mg(N)·kg−1。同列不同小写字母表示同一时间不同处理间在P<0.05水平差异显著。ns表示未达到显著水平, *代表P<0.05显著水平, **代表P<0.01极显著水平。W1: relative water content of 45%±5%; W2: relative water content of 70%±5%; N0: no nitrogen fertilizer; N1: nitrogen application rate of 69 mg(N)·kg−1; N2: nitrogen application rate of 138 mg(N)·kg−1. Different lowercase letters in the same column indicate significant differences at P<0.05 level among treatments in the same date. ns means no significant difference. * means significant differences at P<0.05 level. ** means significant differences at P<0.01 level.
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表 6 干旱胁迫和氮素调控对棉花过氧化氢酶(CAT)活性的影响
Table 6. Effects of water and nitrogen regulation on catalase (CAT) activity of cotton
U·mg−1(protein) 处理
Treatment处理后天数 Days after treatment (d) 0 15 30 45 60 75 90 W1N0 56.08±7.43a 72.58±4.71bc 82.95±8.25bc 93.79±2.27d 87.66±3.74bc 78.94±5.35bc 70.46±4.26d W2N0 57.97±6.74a 65.75±5.10c 74.94±6.16c 84.13±4.90e 77.53±6.01c 67.40±7.15c 62.68±5.21d W1N1 60.56±7.15a 78.71±4.26ab 89.08±6.74b 100.15±1.47bc 95.44±3.24b 85.31±5.89b 78.94±6.10bc W2N1 60.09±12.86a 73.05±5.31bc 81.77±3.56bc 91.67±5.31cd 86.72±4.02bc 76.11±2.48b 69.28±3.74cd W1N2 64.57±3.56a 86.01±2.68a 100.15±3.63a 110.99±4.64a 102.98±8.84a 94.02±2.12a 88.13±5.35a W2N2 65.98±4.71a 80.12±7.63ab 90.02±2.16b 101.57±3.56b 91.20±2.45ab 83.89±4.55ab 77.76±1.87b 水分 Water (W) ns * ** ** ** ** ** 施氮 Nitrogen application (N) ns ** ** ** ** ** ** 水分×氮肥 W×N ns ns ns ns ns ns ns W1: 相对含水量为45%±5%; W2: 相对含水量为70%±5%; N0: 不施氮肥; N1: 施氮69 mg(N)·kg−1; N2: 施氮138 mg(N)·kg−1。同列不同小写字母表示同一时间不同处理间在P<0.05水平差异显著。ns表示未达到显著水平, *代表P<0.05显著水平, **代表P<0.01极显著水平。W1: relative water content of 45%±5%; W2: relative water content of 70%±5%; N0: no nitrogen fertilizer; N1: nitrogen application rate of 69 mg(N)·kg−1; N2: nitrogen application rate of 138 mg(N)·kg−1. Different lowercase letters in the same column indicate significant differences at P<0.05 level among treatments in the same date. ns means no significant difference. * means significant differences at P<0.05 level. ** means significant differences at P<0.01 level.
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表 7 水分和氮素调控对棉花丙二醛(MDA)含量的影响
Table 7. Effects of water and nitrogen regulation on malondialdehyde (MDA) content of cotton
µmol·g−1(FW) 处理
Treatment处理后天数 Days after treatment (d) 0 15 30 45 60 75 90 W1N0 28.68±3.33a 47.65±1.32a 71.91±2.02a 82.69±2.40a 88.64±3.61a 93.46±7.02a 90.38±5.20a W2N0 28.24±2.76b 42.35±2.65b 63.97±2.02b 74.23±1.76b 81.36±2.08b 86.54±5.03ab 82.69±5.45ab W1N1 27.35±2.02bc 41.03±2.65b 61.32±2.02b 70.77±1.76b 78.18±2.08bc 82.31±3.53bc 79.23±2.90bc W2N1 26.91±2.02cd 37.06±2.29c 56.47±0.76c 63.85±2.90c 72.73±2.08cd 76.15±1.15cd 73.46±2.40cd W1N2 25.15±2.65d 36.62±2.76c 54.26±1.32c 60.38±2.90c 68.64±1.57de 73.85±3.05de 70.00±4.37de W2N2 24.71±3.33e 31.32±3.33d 42.35±2.29d 53.08±2.31d 62.73±2.73e 67.31±4.37e 64.62±4.62e 水分 Water (W) ns ** ** ** ** ** ** 施氮 Nitrogen application (N) ns ** ** ** ** ** ** 水分×氮肥 W×N ns ns ns ns ns ns ns W1: 相对含水量为45%±5%; W2: 相对含水量为70%±5%; N0: 不施氮肥; N1: 施氮69 mg(N)·kg−1; N2: 施氮138 mg(N)·kg−1。同列不同小写字母表示同一时间不同处理间在P<0.05水平差异显著。ns表示未达到显著水平, *代表P<0.05显著水平, **代表P<0.01极显著水平。W1: relative water content of 45%±5%; W2: relative water content of 70%±5%; N0: no nitrogen fertilizer; N1: nitrogen application rate of 69 mg(N)·kg−1; N2: nitrogen application rate of 138 mg(N)·kg−1. Different lowercase letters in the same column indicate significant differences at P<0.05 level among treatments in the same date. ns means no significant difference. * means significant differences at P<0.05 level. ** means significant differences at P<0.01 level.
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