Regulating Effect of Exogenous Melatonin on Aluminum Toxicity in Solanum lycopersicum L.
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摘要:
目的 研究褪黑素对铝胁迫下番茄生理上的调控作用，为缓解土壤酸铝化对番茄生长造成的不利影响提供理论依据。 方法 以耐铝品种千禧番茄和铝敏感品种普罗旺斯番茄为供试植物，设置空白组、单铝组、褪黑素（MT）处理组和铝+褪黑素共同施加组，测定各处理组生长周期（10 d、20 d、30 d）过程中番茄叶片抗氧化酶[超氧化物歧化酶（SOD）、过氧化物酶（POD）和过氧化氢酶（CAT）]活性、丙二醛（MDA）、脯氨酸（Pro）、抗坏血酸（AsA）、谷胱甘肽（GSH）含量、叶绿素含量和叶绿素荧光参数，检测根部铝富集含量和根系活力，并对番茄的根、茎、叶进行DNA损伤检测。 结果 铝逆境下两品种番茄生长发育均受到严重抑制，各器官DNA均受损严重，且其根系DNA损伤最为严重。施用褪黑素能够明显缓解番茄的铝毒症状，两品种番茄株高、根长及叶面积等主要生长指标均有所增长；抗氧化酶活性得到显著提高，且随实验周期延长呈现先上升后下降的趋势；GSH和AsA含量最大增幅达91.14%、13.52%；MDA含量降幅最大可达38.39%，脯氨酸含量增幅可达144.81%，调节番茄体内渗透平衡；叶绿素含量明显升高，各项荧光参数得到改善，恢复其光合能力；千禧番茄和普罗旺斯番茄根系活力增幅分别为5.19%和43.03%，两者的根、茎、叶铝富集能力均降低。同时，施加褪黑素也能够有效减缓番茄各器官DNA拖尾现象，修复细胞DNA损伤。 结论 施加外源褪黑素能有效激活铝胁迫下番茄的各项生理响应，促进其生长发育，不仅能提高抗氧化酶系统活性，增强光合能力与根系活力，还可以减少植株体内铝含量，降低细胞DNA受损程度，进一步提升其抗铝能力，为探索MT对逆境下植物的调控作用提供新思路。 Abstract:Objective Regulating effect of melatonin application on the physiology of a tomato plant under aluminum stress was studied. Method Al-tolerant Millennial tomato and Al-sensitive Provence tomato plants were grown in blank control, Al-added, melatonin-added (MT), or Al-and-melatonin-added (AMT) potting soils. Activities of superoxide dismutase (SOD), peroxidase (POD), and catalase (CAT), contents of malondialdehyde (MDA), proline (Pro), ascorbic acid (AsA), glutathione (GSH), and chlorophyll as well as chlorophyll fluorescence parameters, Al-accumulation, and root activity in the plants were monitored. DNA damage occurred to the roots, stems, and leaves of the plants during the experimentation were recorded. Statistical analysis was performed on all collected data using the one-way ANOVA and Duncan test. Result Under Al-stress, the growth and development of the tomato plants of either variety were severely inhibited, and the DNA in the organs, especially the roots, seriously damaged. Whereas the presence of melatonin in the soil (MT and/or AMT ) significantly alleviated the symptoms of Al-toxicity on the plants. For instance, with the addition, the main growth indicators, such as plant height, root length, and leaf area, were improved; the activity of antioxidant enzymes significantly heightened with an increasing trend initially and followed by a decline as treatment duration prolonged; the contents of GSH and AsA raised by 91.14% and 13.52% respectively; the reduction on MDA maximized at 38.39%; proline, which regulates the osmotic balance in tomato, increased to 144.81% and chlorophyll significantly; the fluorescence parameters improved; the photosynthetic capacity much restored; the root activity of Millennial tomato plant increased by 5.19%, and that of Provence tomato by 43.03%; the Al-accumulation in the roots, stems, and leaves significantly reduced; the DNA tailing in organs slowed; and the repairs on damaged DNA in cells observed. Conclusion Applying exogenous melatonin in soil effectively activated various physiological responses, raised the activity of antioxidant enzymes, enhanced the plant photosynthesis, reduced damage on cellular DNA, and elevated the resistance to Al-toxicity of the tomato plants under the heavy metal stress. -
Key words:
- Melatonin /
- Solanum lycopersicum /
- aluminum stress /
- physiological response /
- DNA damage
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图 1 各处理下番茄根系活力的变化
不同小写字母表示同一品种不同处理间差异显著（P＜0.05）。
Figure 1. Changes on tomato root activity under treatments
Datas with different lowercase letters indicate significant differences between treatments on same tomato variety at P＜0.05.
图 2 外源MT对铝胁迫下番茄DNA损伤的影响
Figure 2. Effect of MT on DNA damage on tomato plants under Al-stress
表 1 外源MT对铝胁迫下番茄生长特性的影响
Table 1. Effect of MT on growth of tomato plants under Al-stress
品种
Varieties处理组
Treatments株高
Plant height/cm叶面积
leaf area/cm2根长
Root length/cm千禧 Qianxi CK 64.25±1.35 b 49.16±2.60 a 9.75±0.76 a Al 59.32±1.44 c 33.09±2.24 b 4.33±2.35 b MT 69.16±2.07 a 60.76±3.14 a 11.59±1.35 a Al+MT 65.92±3.12 b 55.40±1.47 a 10.47±1.32 a 普罗旺斯 Puluowangsi CK 59.37±1.21 b 37.95±1.24 a 12.64±1.84 a Al 51.31±2.74 c 25.88±3.27 b 8.24±2.46 b MT 62.81±2.68 a 41.84±1.55 a 16.05±1.27 a Al+MT 60.44±1.67 ab 39.62±2.47 a 14.18±0.55 a 表内数据为平均值±标准误差，同项同列数据后相同字母表示无显著差异（P > 0.05），下同。
Datas are mean ± standard error; those with same letter on same line indicate no significant difference ( P>0.05). Same for below. 
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表 2 外源MT对铝胁迫下番茄抗氧化酶的影响
Table 2. Effects of MT on antioxidant enzymes of tomato plants under Al-stress
测量指标
Measured indicators处理
Treatment千禧 Qianxi 普罗旺斯 Puluowangsi 10 d 20 d 30 d 10 d 20 d 30 d SOD/(U·g−1) CK 3902.26±334.93 b 950.64±36.19 c 675.79±15.47 a 4457.17±75.413 a 906.62±17.75 d 583.64±12.05 b AI 6536.58±237.21 a 1288.10±16.24 ab 529.74±5.01 b 5609.25±138.00 a 1139.10±25.28 b 472.19±7.09 c MT 4421.31±299.02 b 1142.93±41.86 bc 694.85±33.19 a 4712.67±180.63 a 1041.19±24.62 c 595.65±29.14 b AI+MT 8008.05±193.70 a 1407.24±34.62 a 629.48±11.85 a 6600.04±281.52 a 1314.23±32.45 a 813.36±28.87 a POD/
(△OD470·min−1·g−1)CK 58.25±2.72 a 75.33±2.21 b 163.72±7.11 c 110.56±12.98 bc 113.50±8.86 b 118.62±10.85 c AI 55.01±14.66 c 95.76±11.25 b 97.96±13.18 d 42.86±8.97 bc 138.95±9.53 ab 95.42±4.70 d MT 64.88±7.39 ab 80.43±11.95 b 192.96±3.54 b 93.16±6.10 bc 124.86±8.80 ab 171.23±6.18 b AI+MT 73.31±3.57 a 184.41±26.62 a 238.29±9.09 a 114.05±7.30 ab 152.73±5.39 a 193.36±10.77 a CAT/
(μmol·min−1·g−1)CK 5.89±1.53 c 18.36±3.04 b 58.60±1.42 b 9.06±0.99 d 18.50±3.72 d 73.77±1.31 b AI 24.54±3.74 b 32.18±2.51 b 53.00±0.98 c 69.52±2.06 c 60.60±0.92 c 52.53±1.32 c MT 12.03±3.40 c 21.47±4.46 b 29.31±1.43 b 63.31±1.43 b 73.77±0.49 b 83.63±1.89 a AI+MT 61.12±1.96 a 77.17±1.21 a 74.67±1.39 a 74.67±1.39 a 89.77±2.03 a 82.17±1.88 a
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表 3 外源MT对铝胁迫下番茄内源性抗氧化物的影响
Table 3. Effect of MT on antioxidants in tomato plants under Al-stress
测量指标
Measured indicators处理
Treatment千禧 Qianxi 普罗旺斯 Puluowangsi 10 d 20 d 30 d 10 d 20 d 30 d GSH含量
GSH content/(μmol·g−1)CK 0.44±0.01 d 1.07±0.043 a 1.06±0.05 c 0.66±0.01 d 1.11±0.02 b 1.10±0.02 b AI 1.14±0.01 b 0.86±0.03 a 0.89±0.03 d 1.09±0.02 b 1.06±0.02 c 0.66±0.02 c MT 0.61±0.01 c 1.20±0.02 a 1.69±0.02 a 0.71±0.01 c 1.26±0.01 b 1.30±0.04 a AI+MT 1.20±0.02 a 1.65±0.09 a 1.49±0.05 b 1.13±0.02 a 1.57±0.01 a 1.39±0.02 a AsA含量
AsA content/(mg·g−1)CK 0.03±0.01 d 0.18±0.02 c 0.22±0.03 b 0.14±0.01 d 0.12±0.01 d 0.20±0.02 b AI 0.36±0.01 b 0.39±0.01 b 0.11±0.02 c 0.30±0.05 b 0.28±0.05 b 0.13±0.02 c MT 0.36±0.01 c 0.41±0.02 ab 0.37±0.02 a 0.16±0.03 c 0.16±0.04 c 0.22±0.02 b AI+MT 0.40±0.02 a 0.44±0.02 a 0.36±0.03 a 0.31±0.03 a 0.29±0.03 a 0.26±0.04 a
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表 4 外源MT对铝胁迫下番茄脯氨酸和MDA含量的影响
Table 4. Effects of MT on Pro and MDA contents in tomato plants under Al-stress
测量指标
Measured indicators处理
Treatment千禧 Qianxi 普罗旺斯 Puluowangsi 10 d 20 d 30 d 10 d 20 d 30 d MDA含量
MDA content/(nmol·g−1)CK 8.08±0.71 b 6.98±0.40 b 6.19±0.49 c 6.41±0.24 b 6.27±0.72 ab 7.77±0.65 c AI 9.47±0.65 a 8.03±0.47 a 8.01±0.48 a 7.49±0.73 a 7.11±0.70 a 10.17±0.93 a MT 3.43±0.95 ab 4.12±1.15 b 5.83±0.68 c 2.36±0.46 c 5.49±0.52 b 6.95±0.80 c AI+MT 8.75±0.63 c 6.76±0.65 c 6.82±0.62 b 7.20±0.22 a 6.52±0.85 ab 8.72±0.38 b 脯氨酸含量
Proline content/(mg·g−1)CK 4.94±1.33 c 6.39±1.16 c 16.47±1.90 d 2.24±0.97 c 5.10±1.91 b 13.12±1.20 b AI 5.54±1.63 c 9.63±1.43 b 20.23±1.20 c 3.77±0.99 c 8.26±1.82 b 14.43±2.09 b MT 9.64±1.15 b 14.39±1.23 a 34.06±1.06 b 6.48±1.54 b 12.34±2.00 a 25.86±2.03 a AI+MT 12.64±1.20 a 15.34±2.21 a 45.53±2.08 a 13.25±2.86 a 13.89±3.00 a 28.16±3.06 a
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表 5 外源MT对铝胁迫下番茄光合特性的影响
Table 5. Effects of MT on photosynthetic characteristics of tomato plants under Al-stress
测量指标
Measured indicators处理
Treatment千禧 Qianxi 普罗旺斯 Puluowangsi 10 d 20 d 30 d 10 d 20 d 30 d SPAD值
SPAD valueCK 28.17±0.37 a 28.60±0.12 b 35.30±0.32 b 31.37±0.62 b 35.87±0.47 b 35.00±0.38 b AI 24.97±0.32 b 25.67±0.26 c 27.30±0.32 c 28.93±0.77 b 29.67±0.23 c 30.27±0.43 c MT 28.23±0.38 a 30.43±0.32 a 37.03±0.08 a 36.10±0.98 a 40.37±0.27 a 38.73±0.68 a AI+MT 27.03±0.46 a 28.93±0.20 b 34.53±0.55 b 31.40±1.36 b 36.27±0.45 b 34.40±0.10 b 初始荧光
FoCK 0.10±0.03 a 0.11±0.04 b 0.11±0.02 c 0.10±0.03 b 0.10±0.07 d 0.11±0.06 c AI 0.09±0.01 a 0.13±0.04 a 0.13±0.07 a 0.12±0.04 a 0.16±0.01 a 0.18±0.05 a MT 0.06±0.03 c 0.07±0.06 d 0.09±0.05 d 0.09±0.06 c 0.11±0.07 c 0.11±0.04 c AI+MT 0.08±0.02 b 0.09±0.05 c 0.11±0.05 b 0.11±0.03 b 0.12±0.04 b 0.14±0.03 b PSII 最大光化学量子产量
Fv /F mCK 0.92±0.03 b 0.93±0.02 a 0.93±0.04 a 0.82±0.03 d 0.81±0.02 b 0.82±0.03 b AI 0.84±0.02 d 0.73±0.03 d 0.69±0.02 c 0.72±0.03 b 0.65±0.03 d 0.61±0.02 d MT 1.04±0.01 a 1.00±0.01 a 0.94±0.02 a 0.94±0.02 a 0.93±0.03 a 0.95±0.03 a AI+MT 0.88±0.01 c 0.78±0.03 c 0.75±0.01 b 0.79±0.01 c 0.71±0.01 c 0.70±0.02 c
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表 6 外源MT对铝胁迫下番茄铝富集效能的影响
Table 6. Effect of MT on Al-accumulation of tomato plants under Al-stress
品种 处理 Al含量 Al content/(mg·kg-1) Al富集系数 Al enrichment factor 转运系数
Coefficient of transshipment根
Root茎
Stem叶
Leaf土壤
Soil根
Root茎
Stem叶
Leaf千禧 Qianxi CK 0.13±0.00 c 0.10±0.00 b 0.07±0.01 c 0.83±0.03 bc 0.16±0.01 b 0.12±0.01 b 0.08±0.01 ab 1.29±0.005 a Al 0.22±0.02 a 0.14±0.00 a 0.10±0.02 ab 1.02±0.01 a 0.21±0.02 a 0.14±0.01 a 0.10±0.01 a 1.16±0.003 b MT 0.12±0.02 c 0.07±0.01 b 0.06±0.01 b 0.86±0.02 c 0.15±0.02 b 0.09±0.01 c 0.07±0.02 b 1.09±0.002 c Al+MT 0.18±0.01 b 0.13±0.01 a 0.09±0.01 ab 1.11±0.03 ab 0.17±0.01 b 0.12±0.02 b 0.08±0.01 b 1.20±0.002 b 普罗旺斯 Puluo wangsi CK 0.11±0.01 b 0.09±0.01 b 0.05±0.01 b 0.82±0.06 c 0.13±0.01 a 0.11±0.02 ab 0.06±0.01 b 1.37±0.003 b Al 0.15±0.01 a 0.13±0.01 a 0.09±0.001 a 1.02±0.01 ab 0.15±0.03 a 0.13±0.01 a 0.09±0.01 a 1.47±0.004 a MT 0.10±0.01 b 0.08±0.01 b 0.05±0.006 b 0.85±0.03 bc 0.12±0.01 a 0.10±0.01 b 0.06±0.01 b 1.38±0.002 b Al+MT 0.14±0.01 a 0.12±0.01 a 0.08±0.002 a 1.10±0.01 a 0.14±0.01 a 0.11±0.01 b 0.07±0.01 ab 1.32±0.001 c
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