牟萌1,
李保华1,
董学会2,,
1.潍坊市农业科学院 潍坊 261071
2.中国农业大学农学院 北京 100193
基金项目: 国家“重大新药创制”科技重大专项20122X09304006
详细信息
作者简介:牛晓雪, 主要从事种子生理研究。E-mail:apple.xiaoxue@163.com
通讯作者:董学会, 主要研究方向为种子生理学。E-mail:xuehuidong@cau.edu.cn
中图分类号:S330.2计量
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出版历程
收稿日期:2018-03-13
录用日期:2018-06-30
刊出日期:2018-12-01
Physiological mechanism of FeSO4 priming improvement of seed germination performances of Gentiana macrophylla
NIU Xiaoxue1, 2,,MU Meng1,
LI Baohua1,
DONG Xuehui2,,
1. Weifang Academy of Agricultural Sciences, Weifang 261071, China
2. College of Agronomy and Biotechnology, China Agricultural University, Beijing 100193, China
Funds: the National Science and Technology Project of "Major New Drug Creation" of China20122X09304006
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Corresponding author:DONG Xuehui, E-mail: xuehuidong@cau.edu.cn
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摘要
摘要:为探讨FeSO4引发提高秦艽种子萌发抗性的生理机制，同时设置水引发处理，以未作任何处理的种子为对照（CK），研究了0.6% FeSO4引发处理24 h对种子膜透性、储藏性物质、能量、激素及抗性相关酶等方面的影响。结果表明：FeSO4引发降低了种子膜透性，提高种子活力，在吸水12 h和24 h时，电导率分别较CK降低了6.61%和11.67%；调动了种子内部储藏物质的代谢，蔗糖含量减少9.57%，可溶性蛋白增加49.63%，饱和脂肪酸中的豆蔻酸和木焦油酸甲酯含量分别增加4.93%和9.03%，不饱和脂肪酸中的山嵛酸、油酸、亚油酸和亚麻酸分别减少6.73%、8.18%、8.40%和6.70%；改变了种子激素平衡，其中脱落酸的含量下降64.78%，赤霉素含量增加近22倍；加速了种子能量代谢，ATP含量增加2.16倍，细胞色素C氧化酶活性增加67.91%。此外，FeSO4引发使线粒体非酶促系统中谷胱甘肽含量和抗坏血酸的含量分别增加74.08%和10.89%，使细胞内酶促系统超氧化物歧化酶、过氧化氢酶和抗坏血酸过氧化物酶活性分别增加285%、179%以及19.6%。因此，FeSO4引发对种子萌发是一个综合过程，一方面可促进种子物质代谢和提高能荷水平，一方面可改善种子内部生理状态和提高胁迫响应能力，从而提高种子活力，促进种子快速和整齐萌发。
关键词:种子引发/
秦艽/
FeSO4引发/
水引发/
萌发/
生理机制
Abstract:Gentiana macrophylla is a widely used ingredient in traditional Chinese medicine for more than 2 000 years.Now, it is under third-class protection in China and on the list of National Key Protected Wild Herbs.G.macrophylla is a seed-propagated plant with extremely low germination rate.Short seed vigor, lower seed yield and after-ripening severely hinder its seedling establishment and significantly raise nursery cost.Various priming methods have shown that ferrous priming is better than other liquid priming, having higher germination performance under both optimal and adverse environments than non-primed seeds.The objectives of this study was to investigate the physiological mechanism of FeSO4 priming increasing seed germination performance of G.macrophylla to lay the theoretical basis for FeSO4 priming in abiotic stress response and application of FeSO4 priming technology in other plants seeds.Compare with non-priming of seeds, we investigated the effects of 0.6% FeSO4 priming on seed membrane permeability, stored material metabolism, energy metabolism, hormone homeostasis and enzyme activity.In this study, we noted that FeSO4 reduced seed membrane permeability and improved seed vigor by decreasing electrical conductivity by 6.61% at 12 h, 11.67% at 24 h.The mobilization of stored materials such as sugars, proteins and fatty acids were activated, sucrose content decreased by 9.57%, and soluble protein content increased by 49.63%.With regard fatty acids, content of saturated fatty acids increased by 4.93% for myristic acid and 9.03% for methyl lignocerate.Then the content of unsaturated fatty acids decreased by 6.73% for behenic acid, 8.18% for oleic acid, 8.40% for linoleic acid and 6.70% for inolenic acid.In addition, FeSO4 priming altered hormone homeostasis of seeds between ABA and GA, of which ABA content decreased by 64.78% and GA content increased by nearly 22 times.In terms of energy metabolism, FeSO4 priming remarkably improved energy level, increased ATP content by 2.16 times and increased cytochrome C oxidase activity by 67.91%.Moreover, FeSO4 priming acted as an initial stress-exposure, both enzyme-catabolized and non-enzyme-catabolized systems response to environmental stress were activated.This was specifically so for the activities of SOD, CAT, APX of enzyme-catabolized elements, which increased by 2.85 times, 1.79 times and 19.6%, respectively.The contents of non-enzyme-catabolized elements were increased by 74.08% for glutathione and 10.89% for ascorbic acid.In summary, our study indicated that regulation of FeSO4 priming was a complex process that promoted stored material mobilization, ATP biosynthesis, key enzyme activation, phytohormone homeostasis between ABA and GA; and simultaneously regulated both enzymatic-catabolized and non-enzymatic-catabolized effecters in biotic and an-biotic stress.On the one hand, priming induced more advanced development in seeds towards complete germination than unprimed ones.On the other hand, priming was considered as a pre-germination stress-exposure that left seeds with "stress-memory" and improved anti-adversity ability.These changes were not eliminated even when seeds dried to original moisture content.This was the main reason why priming enhanced seed vigor and rapid, uniform seed germination performance.
Key words:Seed priming/
Gentiana mcrophylla Pall./
Ferrous sulfate priming/
Hydro-priming/
Germination/
Physiological mechanism
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图1不同引发剂处理对秦艽种子电导率的影响
同一时间不同字母表示处理间差异显著(P < 0.05)。At the same soaking time, different lowercase letters indicate significant differences among different treatments (P < 0.05).
Figure1.Effect of seed priming with different agents on electrical conductivity of Gentiana macrophylla seeds
下载: 全尺寸图片幻灯片表1不同引发剂处理对秦艽种子内储藏物质的影响
Table1.Effect of seed priming with different agents on storage materials metabolism of Gentiana macrophylla seeds
| mg·g-1 | ||||||
| 引发剂 Priming agent | 葡萄糖 Glucose | 蔗糖 Sucrose | 果糖 Fructose | 淀粉 Starch | 可溶性总糖 Soluble sugar | 可溶性蛋白 Soluble protein |
| CK | 8.23±0.65ab | 56.37±1.41a | 36.07±0.76a | 248.11±2.99a | 95.82±3.57a | 34.01±2.39c |
| H2O | 8.27±0.55a | 56.17±1.39a | 36.25±0.92a | 243.97±1.42a | 94.36±0.66a | 38.43±1.84b |
| FeSO4 | 7.18±0.53b | 51.01±2.36b | 35.43±1.19a | 246.29±5.00a | 94.06±0.90a | 50.89±1.99a |
| 同列不同字母表示差异显著(P < 0.05)。Different lowercase letters in the same column indicate significant differences (P < 0.05). | ||||||
下载: 导出CSV表2不同引发剂处理对秦艽种子脂肪酸含量的影响
Table2.Effect of seed priming with different agents on fatty acids contents of Gentiana macrophylla seeds
| mg·g-1 | |||
| 脂肪酸种类Fatty acid | CK | H2O | FeSO4 |
| 棕榈酸Palmitic acid (C16_0) | 1.001 5±0.039 9a | 0.981 1±0.025 6a | 0.948 9±0.022 5a |
| 硬脂酸Stearic acid (C18_0) | 0.475 2±0.002 8a | 0.451 0±0.008 1b | 0.461 9±0.015 6ab |
| 花生一烯酸Arachidonic acid(C20_0) | 0.054 4±0.002 1a | 0.054 6±0.001 0a | 0.055 1±0.001 9a |
| 山嵛酸甲酯Methyl behenate (C22_0) | 0.024 6±0.002 5a | 0.027 2±0.000 7a | 0.026 6±0.001 2a |
| 豆蔻酸Myristic acid (C14_0) | 0.009 7±0.000 3b | 0.009 7±0.000 1b | 0.010 2±0.000 1a |
| 木焦油酸甲酯Methyl Lignocerate (C24_0) | 0.031 0±0.001 0b | 0.033 1±0.001 1ab | 0.033 8±0.001 4a |
| 棕榈一烯酸Palmitoleic acid (C16_1) | 0.024 2±0.000 9a | 0.023 7±0.000 7a | 0.022 8±0.001 0a |
| 山嵛酸Behenic acid (C20_1) | 0.053 5±0.000 7a | 0.052 1±0.001 6ab | 0.049 9±0.001 8b |
| 油酸Oleic acid (C18_1) | 5.009 8±0.059 9a | 4.869 6±0.054 4a | 4.602 8±0.199 7b |
| 亚油酸Linoleic acid (C18_2) | 11.878 1±0.097 3a | 11.463 0±0.310 5ab | 10.857 3±0.591 2b |
| 亚麻酸Linolenic acid (C18_3) | 0.209 4±0.002 9a | 0.202 8±0.005 4ab | 0.195 2±0.011 1b |
| 同行不同字母表示差异显著(P < 0.05)。Different lowercase letters in the same line indicate significant differences (P < 0.05). | |||
下载: 导出CSV表3不同引发剂处理对秦艽种子激素(ABA、GA)含量、GA/ABA比值及ATP含量的影响
Table3.Effect of seed priming with different agents on hormone contents (ABA, GA), ratio of GA/ABA and ATP content of Gentiana macrophylla seeds
| 引发剂 Priming agent | 脱落酸(ABA) Abscisic acid (μg·g-1) | 赤霉素(GA) Gibberellin (μg·g-1) | GA/ABA比值 GA/ABA ratio | 腺苷三磷酸(ATP) Adenosine triphosphate (μmol·g-1) |
| CK | 4.45±0.34a | 0.16±0.03c | 0.06±0.01c | 9.08E-6±1.59E-7b |
| H2O | 1.89±0.08b | 0.21±0.01b | 0.09±0.02b | 9.04E-6±2.93E-7b |
| FeSO4 | 1.57±0.06c | 3.51±0.23a | 2.26±0.24a | 2.87E-5±2.50E-6a |
| 同列中不同字母表示差异显著(P < 0.05)。Different lowercase letters in the same column indicate significant differences (P < 0.05). | ||||
下载: 导出CSV表4不同引发剂处理对秦艽种子线粒体完整性的影响
Table4.Effect of seed priming with different agents on mitochondrial integrity of Gentiana macrophylla seeds
| 引发剂 Priming agent | 细胞色素C氧化酶(COX) Cytochrome C oxidase (U·g-1·min-1) | 线粒体完整性 Mitochondrial integrity (%) | ||
| 加入Triton前Before adding triton | 加入Triton后After adding triton | |||
| CK | 0.458±0.002a | 0.508±0.015a | 9.873±0.438b | |
| H2O | 0.573±0.018b | 0.637±0.028b | 8.682±2.219ab | |
| FeSO4 | 0.797±0.018c | 0.853±0.039c | 5.565±1.476a | |
| 同列不同字母表示差异显著(P < 0.05)。Different lowercase letters in the same column indicate significant differences (P < 0.05). | ||||
下载: 导出CSV表5不同引发剂对秦艽种子线粒体内非酶促保护系统酶的影响
Table5.Effects of seed priming with different agents on non-enzymatic antioxidants in mitochondria of Gentiana macrophylla seeds
| 引发剂 Priming agent | 还原性谷胱甘肽 (GSH) Reduced glutathione [μmol·g-1(protein)] | 氧化性谷胱甘肽(GSSH) Oxidized glutathione [μmol·g-1(protein)] | 比值 Ratio (GSH/GSSH) | 抗坏血酸(ASC) Ascorbic acid [μmol·g-1(protein)] | 脱氢抗坏血酸(DHA) Dehydroascorbic acid [μmol·g-1(protein)] | 比值 Ratio (ASC/DHA) |
| CK | 23.88±0.86c | 83.71±0.45a | 0.31±0.04c | 12.95±0.45b | 3.67±0.81b | 3.68±1.04a |
| H2O | 27.74±1.98b | 67.26±0.42b | 0.41±0.03b | 15.08±0.40a | 6.15±0.99a | 2.49±0.38ab |
| FeSO4 | 41.57±5.54a | 64.80±4.77b | 0.64±0.08a | 14.36±0.87a | 7.90±2.17a | 1.89±0.38b |
| 同列不同字母表示差异显著(P < 0.05)。Different lowercase letters in the same column indicate significant differences (P < 0.05). | ||||||
下载: 导出CSV表6不同引发剂对秦艽种子抗性相关酶的影响
Table6.Effect of seed priming with different agents on resistance-related enzymes activities of Gentiana macrophylla seeds
| 引发剂 Priming agent | 过氧化氢酶活性 Catalase activity (U·mg-1·min-1) | 抗坏血酸过氧化物酶活性 Ascorbate peroxidase activity (U·mg-1·min-1) | 超氧化物歧化酶活性 Superoxide dismutase activity (mg·g-1) | 丙二醛含量 Malondialdehyde content (mg·g-1) |
| CK | 0.82±0.29c | 0.72±0.23c | 1.43±0.02b | 3.23±0.19b |
| H2O | 2.07±0.63b | 1.48±0.23b | 1.46±0.01b | 2.73±0.19c |
| FeSO4 | 3.17±0.28a | 2.02±0.17a | 1.70±0.03a | 5.97±0.32a |
| 同列不同字母表示差异显著(P < 0.05)。Different lowercase letters in the same column indicate significant differences (P < 0.05). | ||||
下载: 导出CSV参考文献
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