H_2O_2对甘薯幼苗不定根的生长和植株抗冷性的影响
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摘要
本研究针对甘薯在北方推广种植中常遇到的苗期干旱导致存活率低以及常遭受冷害这两个关键性问题,我们以前人的关于H_2O_2在植株根系生长和抗逆性的方面以及转入抗氧化酶基因在植株抗逆性的作用为指导,用转Cu/Zn SOD和APX基因甘薯幼苗以及其非转基因对照植株为实验材料,采用水培和盆栽相结合的方法,研究了外源H_2O_2对幼苗进行根部处理能否促进甘薯幼苗不定根的生长以及同时运用外源H_2O_2对甘薯幼苗盆栽植株进行叶片喷施处理和转入抗氧化酶基因这两种策略能否解决甘薯的抗冷性问题,得到如下主要结果:
1.将不同浓度的外源H_2O_2以及其清除剂抗坏血酸(AsA)加入未转基因甘薯幼苗的水培液中进行6天的实验处理。结果显示:
(1)较低浓度的H_2O_2(0.5 mmoL/L和2.5mmoL/L)处理,促进了甘薯幼苗不定根系的生长,处理组每株根系的总重量、根数目、平均根长、总根表面积以及单根活力均显著高于对照,其中0.5mmoL/L处理组诱导作用最为明显;当浓度达到5mmoL/L时,则表现出显著的抑制作用,幼苗不定根系的生长几乎全部受到了抑制,而且幼根还受到了严重的伤害;用4.0 mmol/L和10.0 mmoL/L的AsA处理组也都表现出显著的抑制作用,不但不定根系的形成完全受到抑制,还阻碍了幼苗茎截面处的损伤修复;而先用2.5 mmoL/L H_2O_2处理3天后再用4.0mmoL/L AsA处理3天实验组则表现出一定的根系伸长抑制特性,但根系数目显著增多。
(2)进一步分析叶片损伤程度、光合色素以及内源H_2O_2含量变化情况,发现内源H_2O_2、MDA含量在各个处理之间没有显著差异,但均高于对照组,而光合色素在各个处理组间变化情况与植株的根系生长状况呈现出一致性。
2.以转Cu/Zn SOD和APX基因甘薯苗以及其非转基因对照植株为实验材料,研究了其在5℃冷胁迫一夜(12h)后植株的冷后恢复情况以及用1.0 mmoL/L H_2O_2于冷胁迫前4h预处理对其抗冷性增强的问题。结果显示:
(1)在短时间(12h)冷胁迫后的很短时间(2h)内,非转基因植株表现出APX,SOD,CAT这几种抗氧化酶活性以及类胡萝卜素这一非酶类抗氧化剂含量的显著降低,膜透性显著升高,光合色素以及光合电子传递链受到破坏,光合速率显著下降;在经历一天多的室温恢复后,植株的上述几种抗氧化酶活性以及类胡萝卜素含量都得到显著提升,而膜透性继续增加,光合电子传递链得到完全修复,光合色素却显著下降,但光合作用得到了较大程度的恢复。
(2)在冷胁迫后,转基因植株较对非转基因对照植株上述几种抗氧化酶和类胡萝卜素表现出了更高的活性和含量,较低的膜透性,具有较强的光合色素以及光合电子传递系统的保护能力,因而其光合速率相对较强,显示出了相对较强的抗冷胁迫能力。
(3)用1.0 mmoL/L H_2O_2预处理组的植株较未预处理组在冷胁迫后的恢复过程中,表现出了较强的上述抗氧化酶活性的恢复能力,较低的膜损伤物质MDA的产生量以及较低的膜通透性,且在冷处理后明显表现出较良好的生长状况,因而得出用1.0mmoL/L H_2O_2预处理植株能够显著提高甘薯幼苗的冷后恢复能力,增强其抗冷性。
3.从上述两实验结果的得出:(1)可以通过对幼苗根系进行外源H_2O_2处理来促进甘薯幼苗不定根系的大量形成和快速生长,其处理的最适浓度为0.5mmoL/L;(2)通过转入Cu/Zn SOD和APX基因能够增强甘薯幼苗的抗冷性,且在冷害到来前用1.0mmoL/L H_2O_2预处理植株亦能够使甘薯幼苗的抗冷性得到提高,而同时运用这两种策略则能够更显著地提高植株的抗冷性。
When we try to extend sweetpotato planting area to the north of china,We often faced two main key problems, these are drought-induced low survival rate and confronted with chilling stress.We take the former studies,which on the H_2O_2 in plant root growth and stress resistance as well as the role of transfer antioxidant enzyme gene in plants to stress resistance, as a guide, Using transgenic sweet potato that expressing both Cu/Zn SOD and APX in chloroplasts and its non-transgenic control plant as materials, to study on whether the adding of exogenous H_2O_2 in seedlings culture solution to promote the adventitious roots growth, and also on whether the use of exogenous H_2O_2 sprayed on the leaves of potted sweet potato seedling as well as the transfer antioxidant enzyme gene strategy could resolve these chilling resistance problem of sweetpotato.And then we got these results as following:
1. In the experiment, varies concentrations of exogenous hydrogen peroxide as well as a hydrogen peroxide scavenger ascorbic acid were added into the sweetpotato seedlings’culture solution to study the effect of H_2O_2 on adventitious roots and leaves growth of sweetpotato seedling. Results showed that:
(1) When sweetpotato seedling cultured in lower concentrations H_2O_2 solutions (0.5mmol/L and 2.5 mmol/L) had display an inducement on adventitious root formation and growth, the total root weight, root number, total root length and total root surface area per seedling, as well as root activity of were significantly higher than the control which added nothing, especially in 0.5 mmol/L H_2O_2 treatment; As the concentration of H_2O_2 reached to 5 mmol/L, almost the whole growth of adventitious roots were significant inhibited, roots were seriously damaged also; While added with AsA in the culture solution also showed a significant inhibition on adventitious roots growth, neither the concentration was 4.0 mmol/L nor 10.0 mmol/L, the formation of adventitious roots has been completely inhibited, the damage-reparation on the stem cross-section of the cutting seedlings has also been hindered; However, in the treatment with 4 mmol/L AsA added after treated with 2.5 mmol/L H_2O_2 for three days, the elongation of adventitious roots was inhibited to a certain degree.
(2) It has been found that there has a consistency between the growth status of adventitious roots and the leaves growth status, leaves MDA content, leaves water content as well as the changes in the photosynthetic pigments by further analyze the leaves physiology metabolism status, but no consistency has been discovered between the concentrations of endogenous H_2O_2 in leaves and the exogenous H_2O_2 concentrations which added in culture solution. In conclusion, the adventitious roots growth could be induced by exogenous H_2O_2 below the injury concentration (5.0 mmol/L) and it could be reversed by AsA treatment, it also showed that a certain concentration of accumulated endogenous H_2O_2 is indispensable for the plants in the course of adventitious roots formation. H_2O_2 effect on leaves’physiological process was result from the effects it did on roots growth status first, and then the growth status of roots affect the leaves growth status. The best inducement on adventitious roots and leaves growth was found in 0.5 mmol/L H_2O_2 treatment during our experiment.
2. We take transgenic potted sweetpotato that expressing both Cu/Zn SOD and APX in chloroplasts and its non-transgenic control plant as materials, study on the recoverability of them after one night (12h) chilling stress at 5℃, and also the enhancement of chilling resistance when treated them with 1.0 mmol/L H_2O_2 before chilling stress. The results showed that:
(1) Soon after a short time (12h) chilling stress, the activity of anti-oxidative enzymes as SOD, AXP and CAT in non-transgenic sweet potato decreased significantly and so did the content of carotenoid, but the membrane permeability increased significantly, photosynthetic pigment and photosynthetic electron transport chain were damaged, so the photosynthetic rate reduced; while after one day recovery in room temperature, the activities of above mentioned anti-oxidative enzymes and the content of carotenoid all increased significantly, membrane permeability also increased continuously, and photosynthetic electron transport chain was repaired completely while photosynthetic pigment decreased significantly, but the photosynthesis recovered to a large extent.
(2) After chilling stress, compared with non-transgenic sweet potato, it could be observed that in transgenic sweet potato, the activities of above mentioned anti-oxidative enzymes and the content of caroternoid were higher, membrane permeability was lower, have a stronger ability to protect photosynthetic pigment and photosynthetic electron transport chain, therefore the photosynthetic rate was relatively higher, so a stronger chilling resistance was showed in transgenic sweet potato.
(3) It could also be observed in the recovery process of the plants that had been treated with 1.0 mmol/L H_2O_2 compared with non-treatments before chilling stress, the recovery capabilities of the anti-oxidative enzymes were stronger, the content of MDA and membrane permeability were lower, and the growth status after chilling was better, therefore it could be concluded that the pretreatment with 1.0 mmol/L H_2O_2 could improve the recovery capability of the plant after chilling and enhance the chilling resistance.
3. From results of the above two experiment, we can obtained: (1) When treated root with exogenous H_2O_2, it can promote the formation of sweetpotato seedling adventitious roots to a large number and accelerate root elongation growth, the optimal induce concentration is 0.5mmoL / L; Through transfer the Cu/Zn SOD and APX genes in sweetpotato can enhance its chilling resistance, and sprayed with 1.0mmoL /L H_2O_2 on leaves before the arrival of cold weather could also be able to improve plant chilling resistance , further more,combined theses two strategies would be more significant to improve the cold resistance of plants.
1.将不同浓度的外源H_2O_2以及其清除剂抗坏血酸(AsA)加入未转基因甘薯幼苗的水培液中进行6天的实验处理。结果显示:
(1)较低浓度的H_2O_2(0.5 mmoL/L和2.5mmoL/L)处理,促进了甘薯幼苗不定根系的生长,处理组每株根系的总重量、根数目、平均根长、总根表面积以及单根活力均显著高于对照,其中0.5mmoL/L处理组诱导作用最为明显;当浓度达到5mmoL/L时,则表现出显著的抑制作用,幼苗不定根系的生长几乎全部受到了抑制,而且幼根还受到了严重的伤害;用4.0 mmol/L和10.0 mmoL/L的AsA处理组也都表现出显著的抑制作用,不但不定根系的形成完全受到抑制,还阻碍了幼苗茎截面处的损伤修复;而先用2.5 mmoL/L H_2O_2处理3天后再用4.0mmoL/L AsA处理3天实验组则表现出一定的根系伸长抑制特性,但根系数目显著增多。
(2)进一步分析叶片损伤程度、光合色素以及内源H_2O_2含量变化情况,发现内源H_2O_2、MDA含量在各个处理之间没有显著差异,但均高于对照组,而光合色素在各个处理组间变化情况与植株的根系生长状况呈现出一致性。
2.以转Cu/Zn SOD和APX基因甘薯苗以及其非转基因对照植株为实验材料,研究了其在5℃冷胁迫一夜(12h)后植株的冷后恢复情况以及用1.0 mmoL/L H_2O_2于冷胁迫前4h预处理对其抗冷性增强的问题。结果显示:
(1)在短时间(12h)冷胁迫后的很短时间(2h)内,非转基因植株表现出APX,SOD,CAT这几种抗氧化酶活性以及类胡萝卜素这一非酶类抗氧化剂含量的显著降低,膜透性显著升高,光合色素以及光合电子传递链受到破坏,光合速率显著下降;在经历一天多的室温恢复后,植株的上述几种抗氧化酶活性以及类胡萝卜素含量都得到显著提升,而膜透性继续增加,光合电子传递链得到完全修复,光合色素却显著下降,但光合作用得到了较大程度的恢复。
(2)在冷胁迫后,转基因植株较对非转基因对照植株上述几种抗氧化酶和类胡萝卜素表现出了更高的活性和含量,较低的膜透性,具有较强的光合色素以及光合电子传递系统的保护能力,因而其光合速率相对较强,显示出了相对较强的抗冷胁迫能力。
(3)用1.0 mmoL/L H_2O_2预处理组的植株较未预处理组在冷胁迫后的恢复过程中,表现出了较强的上述抗氧化酶活性的恢复能力,较低的膜损伤物质MDA的产生量以及较低的膜通透性,且在冷处理后明显表现出较良好的生长状况,因而得出用1.0mmoL/L H_2O_2预处理植株能够显著提高甘薯幼苗的冷后恢复能力,增强其抗冷性。
3.从上述两实验结果的得出:(1)可以通过对幼苗根系进行外源H_2O_2处理来促进甘薯幼苗不定根系的大量形成和快速生长,其处理的最适浓度为0.5mmoL/L;(2)通过转入Cu/Zn SOD和APX基因能够增强甘薯幼苗的抗冷性,且在冷害到来前用1.0mmoL/L H_2O_2预处理植株亦能够使甘薯幼苗的抗冷性得到提高,而同时运用这两种策略则能够更显著地提高植株的抗冷性。
When we try to extend sweetpotato planting area to the north of china,We often faced two main key problems, these are drought-induced low survival rate and confronted with chilling stress.We take the former studies,which on the H_2O_2 in plant root growth and stress resistance as well as the role of transfer antioxidant enzyme gene in plants to stress resistance, as a guide, Using transgenic sweet potato that expressing both Cu/Zn SOD and APX in chloroplasts and its non-transgenic control plant as materials, to study on whether the adding of exogenous H_2O_2 in seedlings culture solution to promote the adventitious roots growth, and also on whether the use of exogenous H_2O_2 sprayed on the leaves of potted sweet potato seedling as well as the transfer antioxidant enzyme gene strategy could resolve these chilling resistance problem of sweetpotato.And then we got these results as following:
1. In the experiment, varies concentrations of exogenous hydrogen peroxide as well as a hydrogen peroxide scavenger ascorbic acid were added into the sweetpotato seedlings’culture solution to study the effect of H_2O_2 on adventitious roots and leaves growth of sweetpotato seedling. Results showed that:
(1) When sweetpotato seedling cultured in lower concentrations H_2O_2 solutions (0.5mmol/L and 2.5 mmol/L) had display an inducement on adventitious root formation and growth, the total root weight, root number, total root length and total root surface area per seedling, as well as root activity of were significantly higher than the control which added nothing, especially in 0.5 mmol/L H_2O_2 treatment; As the concentration of H_2O_2 reached to 5 mmol/L, almost the whole growth of adventitious roots were significant inhibited, roots were seriously damaged also; While added with AsA in the culture solution also showed a significant inhibition on adventitious roots growth, neither the concentration was 4.0 mmol/L nor 10.0 mmol/L, the formation of adventitious roots has been completely inhibited, the damage-reparation on the stem cross-section of the cutting seedlings has also been hindered; However, in the treatment with 4 mmol/L AsA added after treated with 2.5 mmol/L H_2O_2 for three days, the elongation of adventitious roots was inhibited to a certain degree.
(2) It has been found that there has a consistency between the growth status of adventitious roots and the leaves growth status, leaves MDA content, leaves water content as well as the changes in the photosynthetic pigments by further analyze the leaves physiology metabolism status, but no consistency has been discovered between the concentrations of endogenous H_2O_2 in leaves and the exogenous H_2O_2 concentrations which added in culture solution. In conclusion, the adventitious roots growth could be induced by exogenous H_2O_2 below the injury concentration (5.0 mmol/L) and it could be reversed by AsA treatment, it also showed that a certain concentration of accumulated endogenous H_2O_2 is indispensable for the plants in the course of adventitious roots formation. H_2O_2 effect on leaves’physiological process was result from the effects it did on roots growth status first, and then the growth status of roots affect the leaves growth status. The best inducement on adventitious roots and leaves growth was found in 0.5 mmol/L H_2O_2 treatment during our experiment.
2. We take transgenic potted sweetpotato that expressing both Cu/Zn SOD and APX in chloroplasts and its non-transgenic control plant as materials, study on the recoverability of them after one night (12h) chilling stress at 5℃, and also the enhancement of chilling resistance when treated them with 1.0 mmol/L H_2O_2 before chilling stress. The results showed that:
(1) Soon after a short time (12h) chilling stress, the activity of anti-oxidative enzymes as SOD, AXP and CAT in non-transgenic sweet potato decreased significantly and so did the content of carotenoid, but the membrane permeability increased significantly, photosynthetic pigment and photosynthetic electron transport chain were damaged, so the photosynthetic rate reduced; while after one day recovery in room temperature, the activities of above mentioned anti-oxidative enzymes and the content of carotenoid all increased significantly, membrane permeability also increased continuously, and photosynthetic electron transport chain was repaired completely while photosynthetic pigment decreased significantly, but the photosynthesis recovered to a large extent.
(2) After chilling stress, compared with non-transgenic sweet potato, it could be observed that in transgenic sweet potato, the activities of above mentioned anti-oxidative enzymes and the content of caroternoid were higher, membrane permeability was lower, have a stronger ability to protect photosynthetic pigment and photosynthetic electron transport chain, therefore the photosynthetic rate was relatively higher, so a stronger chilling resistance was showed in transgenic sweet potato.
(3) It could also be observed in the recovery process of the plants that had been treated with 1.0 mmol/L H_2O_2 compared with non-treatments before chilling stress, the recovery capabilities of the anti-oxidative enzymes were stronger, the content of MDA and membrane permeability were lower, and the growth status after chilling was better, therefore it could be concluded that the pretreatment with 1.0 mmol/L H_2O_2 could improve the recovery capability of the plant after chilling and enhance the chilling resistance.
3. From results of the above two experiment, we can obtained: (1) When treated root with exogenous H_2O_2, it can promote the formation of sweetpotato seedling adventitious roots to a large number and accelerate root elongation growth, the optimal induce concentration is 0.5mmoL / L; Through transfer the Cu/Zn SOD and APX genes in sweetpotato can enhance its chilling resistance, and sprayed with 1.0mmoL /L H_2O_2 on leaves before the arrival of cold weather could also be able to improve plant chilling resistance , further more,combined theses two strategies would be more significant to improve the cold resistance of plants.
引文
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张明生,谢波,戚金亮,谈锋,张启堂,杨永华. 2006.甘薯植株形态,生长势和产量与品种抗旱性的关系.热带作物学报, 27: 39-43
Allan A, Fluhr R. 1997.Two distinct sources of elicited reactive oxygen species in tobacco epidermal cells. The Plant Cell Online, 9: 1559
Allen R, Webb R, Schake S. 1997.Use of transgenic plants to study antioxidant defenses. Free Radical Biology and Medicine, 23: 473-479
Aono M, Kubo A, Saji H, Natori T, Tanaka K, Kondo N. 1991.Resistance to active oxygen toxicity of transgenic Nicotiana tabacum that expresses the gene for glutathione reductase from Escherichia coli. Plant and cell physiology, 32: 691
Bailly C, Benamar A, Corbineau F, Come D. 1996.Changes in malondialdehyde content and in superoxide dismutase, catalase and glutathione reductase activities in sunflower seeds as related to deterioration during accelerated aging. Physiologia Plantarum, 97: 104-110
Bowler C, Slooten L, Vandenbranden S, De Rycke R, Botterman J, Sybesma C, Van Montagu M, Inze D. 1991.Manganese superoxide dismutase can reduce cellular damage mediated by oxygen radicals in transgenic plants. The EMBO Journal, 10: 1723
Bright J, Desikan R, Hancock J, Weir I, Neill S. 2006.ABA-induced NO generation and stomatal closure in Arabidopsis are dependent on H2O2 synthesis. The Plant Journal, 45: 113-122
Delledonne M, Zeier J, Marocco A, Lamb C. 2001.Signal interactions between nitric oxide and reactive oxygen intermediates in the plant hypersensitive disease resistance response. Proceedings of the National Academy of Sciences of the United States of America, 98: 13454
Dunand C, Crèvecoeur M, Penel C. 2007.Distribution of superoxide and hydrogen peroxide in Arabidopsis root and their influence on root development: possible interaction with peroxidases. New Phytologist, 174: 332-341
Foreman J, Demidchik V, Bothwell J, Mylona P, Miedema H, Torres M, Linstead P, Costa S, Brownlee C, Jones J. 2003.Reactive oxygen species produced by NADPH oxidase regulate plant cell growth. Nature, 422: 442-446
Foyer C, Souriau N, Perret S, Lelandais M, Kunert K, Pruvost C, Jouanin L. 1995.Overexpression of glutathione reductase but not glutathione synthetase leads to increases in antioxidant capacity and resistance to photoinhibition in poplar trees. Plant Physiology, 109: 1047
Gupta A, Heinen J, Holaday A, Burke J, Allen R. 1993a.Increased resistance to oxidative stress in transgenic plants that overexpress chloroplastic Cu/Zn superoxide dismutase. Proceedings of the National Academy of Sciences, 90: 1629-1633
Gupta A, Webb R, Holaday A, Allen R (1993b) Overexpression of superoxide dismutase protects plants from oxidative stress (induction of ascorbate peroxidase in superoxide dismutase-overexpressing plants). Am Soc Plant Biol, pp 1067-1073
Henzler T, Steudle E. 2000.Transport and metabolic degradation of hydrogen peroxide in Chara corallina: model calculations and measurements with the pressure probe suggest transport of H2O2 across water channels. Journal of Experimental Botany, 51: 2053
Hodges D, Andrews C, Johnson D, Hamilton R. 1997.Antioxidant enzyme responses to chilling stress in differentially sensitive inbred maize lines. Journal of Experimental Botany, 48: 1105-1113
Joo J, Bae Y, Lee J. 2001.Role of auxin-induced reactive oxygen species in root gravitropism. Science's STKE, 126: 1055
Kim K, Kwon S, Lee H, Hur Y, Bang J, Kwak S. 2003.A novel oxidative stress-inducible peroxidase promoter from sweetpotato: molecular cloning and characterization in transgenic tobacco plants and cultured cells. Plant molecular biology, 51: 831-838
Kratsch H, Wise R. 2000.The ultrastructure of chilling stress. Plant, Cell & Environment, 23: 337-350
Kwon S, Jeong Y, Lee H, Kim J, Cho K, Allen R, Kwak S. 2002.Enhanced tolerances of transgenic tobacco plants expressing both superoxide dismutase and ascorbate peroxidase in chloroplasts against methyl viologen-mediated oxidative stress. Plant, Cell & Environment, 25: 873-882
Lee D, Lee C. 2000.Chilling stress-induced changes of antioxidant enzymes in the leaves of cucumber in gel enzyme activity assay. Plant Science, 159: 75-85
Lee S, Choi H, Suh S, Doo I, Oh K, Jeong Choi E, Schroeder Taylor A, Low P, Lee Y. 1999.Oligogalacturonic acid and chitosan reduce stomatal aperture by inducing the evolution of reactive oxygen species from guard cells of tomato and Commelina communis. Plant Physiology, 121: 147
Li S, Xue L, Xu S, Feng H, An L. 2007.Hydrogen peroxide involvement in formation and development of adventitious roots in cucumber. Plant Growth Regulation, 52: 173-180
Lim S, Kim Y, Kim S, Kwon S, Lee H, Kim J, Cho K, Paek K, Kwak S. 2007.Enhanced tolerance of transgenic sweetpotato plants that express both CuZnSOD and APX in chloroplasts to methyl viologen-mediated oxidative stress and chilling. Molecular Breeding, 19: 227-239
McInnis S, Desikan R, Hancock J, Hiscock S. 2006.Production of reactive oxygen species and reactive nitrogen species by angiosperm stigmas and pollen: potential signalling crosstalk? New Phytologist, 172: 221-228
McKersie B, Bowley S, Harjanto E, Leprince O. 1996.Water-deficit tolerance and field performance of transgenic alfalfa overexpressing superoxide dismutase. Plant Physiology, 111: 1177-1181
McKersie B, Chen Y, De Beus M, Bowley S, Bowler C, Inze D, D'halluin K, Botterman J. 1993.Superoxide dismutase enhances tolerance of freezing stress in transgenic alfalfa (Medicago sativa L.). Plant Physiology, 103: 1155
McKersie B, Murnaghan J, Jones K, Bowley S. 2000.Iron-superoxide dismutase expression in transgenic alfalfa increases winter survival without a detectable increase in photosynthetic oxidative stress tolerance. Plant Physiology, 122: 1427-1438
Mittler R. 2002.Oxidative stress, antioxidants and stress tolerance. Trends in Plant Science, 7: 405-410
Mittler R, Herr E, Orvar B, van Camp W, Willekens H, InzéD, Ellis B. 1999.Transgenic tobacco plants with reduced capability to detoxify reactive oxygen intermediates are hyperresponsive to pathogen infection. Proceedings of the National Academy of Sciences of the United States of America, 96: 14165
Mukherjee S, Choudhuri M. 1983.Determination of glycolate oxidase activity, H2O2 content and catalase activity. Physiol Plant, 58: 167-170
Neill S, Desikan R, Clarke A, Hurst R, Hancock J (2002a) Hydrogen peroxide and nitric oxide as signalling molecules in plants. Soc Experiment Biol, pp 1237-1247
Neill S, Desikan R, Hancock J. 2002b.Hydrogen peroxide signalling. Current Opinion in Plant Biology, 5: 388-395
Perl A, Perl-Treves R, Galili S, Aviv D, Shalgi E, Malkin S, Galun E. 1993.Enhanced oxidative-stress defense in transgenic potato expressing tomato Cu, Zn superoxide dismutases. TAG Theoretical and Applied Genetics, 85: 568-576
Pitcher L, Repetti P, Zilinskas B. 1994.Overproduction of ascorbate peroxidase protects transgenic tobacco against oxidative stress (abstract No. 623). Plant Physiol, 105: 116
Potikha T, Collins C, Johnson D, Delmer D, Levine A. 1999.The involvement of hydrogen peroxide in the differentiation of secondary walls in cotton fibers. Plant Physiology, 119: 849
Potocky M, Jones M, Bezvoda R, Smirnoff N,ársky V. 2007.Reactive oxygen species produced by NADPH oxidase are involved in pollen tube growth. Comparative Biochemistry and Physiology, Part A, 146: 269-270
Prasad T, Anderson M, Martin B, Stewart C. 1994.Evidence for chilling-induced oxidative stress in maize seedlings and a regulatory role for hydrogen peroxide. The Plant Cell Online, 6: 65-74
Ren D, Yang H, Zhang S. 2002.Cell death mediated by MAPK is associated with hydrogen peroxide production in Arabidopsis. Journal of Biological Chemistry, 277: 559-565
Shah K, Kumar R, Verma S, Dubey R.Effect of cadmium on lipid peroxidation, superoxide anion generation and activities of antioxidant enzymes in growing rice seedlings.Время, 22: 16
Slooten L, Capiau K, Van Camp W, Van Montagu M, Sybesma C, InzéD. 1995.Factors affecting the enhancement of oxidative stress tolerance in transgenic tobacco overexpressing manganese superoxide dismutase in the chloroplasts. Plant Physiology, 107: 737
Su G, Zhang W, Liu Y. 2006.Involvement of hydrogen peroxide generated by polyamine oxidative degradation in the development of lateral roots in soybean. Journal of Integrative Plant Biology, 48: 426
Svalheim, Robertsen B. 2006.Elicitation of H2O2 production in cucumber hypocotyl segments by oligo-1, 4-α-D-galacturonides and an oligo-β-glucan preparation from cell walls of Phythophthora megasperma f. sp. glycinea. Physiologia Plantarum, 88: 675-681
Svalheim O, Robertsen B. 1993.Elicitation of hydrogen peroxide production in cucumber hypocotyl segments by oligo-14-alpha-d-galacturonides and an oligo-beta-glucan preparation from cell walls of Phytophthora megasperma f. sp. glycinea. Physiol Plant, 88: 675-681
Tang L, Kwon S, Kim S, Kim J, Choi J, Cho K, Sung C, Kwak S, Lee H. 2006.Enhanced tolerance of transgenic potato plants expressing both superoxide dismutase and ascorbate peroxidase in chloroplasts against oxidative stress and high temperature. Plant Cell Reports, 25: 1380-1386
Tepperman J, Dunsmuir P. 1990.Transformed plants with elevated levels of chloroplastic SOD are not more resistant to superoxide toxicity. Plant molecular biology, 14: 501-511
Tiwari B, Belenghi B, Levine A. 2002.Oxidative stress increased respiration and generation of reactive oxygen species, resulting in ATP depletion, opening of mitochondrial permeability transition, and programmed cell death. Plant Physiology, 128: 1271
Van Camp W, Capiau K, Van Montagu M, Inze D, Slooten L (1996a) Enhancement of oxidative stress tolerance in transgenic tobacco plants overproducing Fe-superoxide dismutase in chloroplasts. Am Soc Plant Biol, pp 1703-1714
Van Camp W, Capiau K, Van Montagu M, Inze D, Slooten L. 1996b.Enhancement of oxidative stress tolerance in transgenic tobacco plants overproducing Fe-superoxide dismutase in chloroplasts. Plant Physiology, 112: 1703
Webb R, Allen R. 1996.Overexpression of pea cytosolic ascorbate peroxidase confers protection against oxidative stress in transgenic Nicotiana tabacum. Plant Physiol. Supplement, 111: 48
YU C, MURPHY T, SUNG W, LIN C. 2002.H2O2 treatment induces glutathione accumulation and chilling tolerance in mung bean. Functional plant biology, 29: 1081-1087
Zhang X, Zhang L, Dong F, Gao J, Galbraith D, Song C. 2001.Hydrogen peroxide is involved in abscisic acid-induced stomatal closure in Vicia faba. Plant Physiology, 126: 1438
白玲,周云,张晓然,宋纯鹏,曹鸣庆. 2007.过氧化氢参与了脱落酸调控的拟南芥根形态发育.科学通报, 52: 608-611
董玉,孟宁,李师翁. 2007.H2O2对黄瓜不定根形成中抗氧化酶活性的影响.兰州交通大学学报, 26: 140-144
简令成,卢存福,李积宏. 2005.适宜低温锻炼提高冷敏感植物玉米和番茄的抗冷性及其生理基础.作物学报, 31: 971-976
李忠光,李江鸿. 2002.在单一提取系统中同时测定五种植物抗氧化酶.云南师范大学学报:自然科学版, 22: 44-48
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李建梅,邓西平. 2007b.干旱和复水条件下转基因甘薯的光合特性.水土保持学报, 4
刘裕强,江玲,孙立宏,王春明,翟虎渠,万建民. 2005.褐飞虱刺吸诱导的水稻一些防御性酶活性的变化.植物生理与分子生物学学报, 31: 643-650
李筠,邓西平,郭尚洙,田中净. 2006.转铜/锌超氧化物歧化酶和抗坏血酸过氧化物酶基因甘薯的耐旱性.植物生理与分子生物学学报, 32: 451-457
李师翁,薛林贵,冯虎元,徐世键,安黎哲. 2007.植物中的H2O2信号及其功能.中国生物化学与分子生物学报, 23: 804-810
逯明辉,宋慧,李晓明,陈劲枫. 2005.冷害过程中黄瓜叶片SOD, CAT和POD活性的变化.西北植物学报, 25: 1570-1573
秦小琼,贾士荣. 1997.植物抗氧化逆境的基因工程(综述).农业生物技术学报, 5: 14-24
王毅,杨宏福. 1994.园艺植物冷害和抗冷性的研究—文献综述.园艺学报, 21: 239-244郁继华,吕军芬,舒英杰. 2004.外源H2O2对西瓜幼苗抗冷性的影响.中国蔬菜: 24-25
邬景禹,孙近友. 1989.我国甘薯种质资源研究的发展战略.中国甘薯, 3: 19-27
许育彬,陈越,付增光. 2004.甘薯的抗旱生理及栽培技术研究进展.干旱地区农业研究, 22: 128-131
许育彬,宋亚珍,李世清. 2009.土壤水分和施肥水平对甘薯叶片气体交换的影响.中国生态农业学报, 17: 79-84
尹永强,胡建斌,邓明军. 2007.植物叶片抗氧化系统及其对逆境胁迫的响应研究进展.中国农学通报, 23: 105-110
吴雨华. 2003.世界甘薯加工利用新趋势.食品研究与开发
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张立明,王庆美,王荫墀. 2003.甘薯的主要营养成分和保健作用.杂粮作物, 23: 162-166
张志良. 2003.过氧化物酶活性的测定.植物生理学实验指南.北京:高等教育出版社: 123—124
张明生,杜建厂,谢波,谈锋,杨永华. 2004a.水分胁迫下甘薯叶片渗透调节物质含量与品种抗旱性的关系.南京农业大学学报, 27: 123-125
张明生,彭忠华,谢波,谈锋,张启堂,付玉凡,杨春贤,杨永华. 2004b.甘薯离体叶片失水速率及渗透调节物质与品种抗旱性的关系.中国农业科学, 37: 152-156
张明生,谈锋. 2003.甘薯可溶性蛋白,叶绿素及ATP含量变化与品种抗旱性关系的研究.中国农业科学, 36: 13-16
张明生,谢波,戚金亮,谈锋,张启堂,杨永华. 2006.甘薯植株形态,生长势和产量与品种抗旱性的关系.热带作物学报, 27: 39-43
Allan A, Fluhr R. 1997.Two distinct sources of elicited reactive oxygen species in tobacco epidermal cells. The Plant Cell Online, 9: 1559
Allen R, Webb R, Schake S. 1997.Use of transgenic plants to study antioxidant defenses. Free Radical Biology and Medicine, 23: 473-479
Aono M, Kubo A, Saji H, Natori T, Tanaka K, Kondo N. 1991.Resistance to active oxygen toxicity of transgenic Nicotiana tabacum that expresses the gene for glutathione reductase from Escherichia coli. Plant and cell physiology, 32: 691
Bailly C, Benamar A, Corbineau F, Come D. 1996.Changes in malondialdehyde content and in superoxide dismutase, catalase and glutathione reductase activities in sunflower seeds as related to deterioration during accelerated aging. Physiologia Plantarum, 97: 104-110
Bowler C, Slooten L, Vandenbranden S, De Rycke R, Botterman J, Sybesma C, Van Montagu M, Inze D. 1991.Manganese superoxide dismutase can reduce cellular damage mediated by oxygen radicals in transgenic plants. The EMBO Journal, 10: 1723
Bright J, Desikan R, Hancock J, Weir I, Neill S. 2006.ABA-induced NO generation and stomatal closure in Arabidopsis are dependent on H2O2 synthesis. The Plant Journal, 45: 113-122
Delledonne M, Zeier J, Marocco A, Lamb C. 2001.Signal interactions between nitric oxide and reactive oxygen intermediates in the plant hypersensitive disease resistance response. Proceedings of the National Academy of Sciences of the United States of America, 98: 13454
Dunand C, Crèvecoeur M, Penel C. 2007.Distribution of superoxide and hydrogen peroxide in Arabidopsis root and their influence on root development: possible interaction with peroxidases. New Phytologist, 174: 332-341
Foreman J, Demidchik V, Bothwell J, Mylona P, Miedema H, Torres M, Linstead P, Costa S, Brownlee C, Jones J. 2003.Reactive oxygen species produced by NADPH oxidase regulate plant cell growth. Nature, 422: 442-446
Foyer C, Souriau N, Perret S, Lelandais M, Kunert K, Pruvost C, Jouanin L. 1995.Overexpression of glutathione reductase but not glutathione synthetase leads to increases in antioxidant capacity and resistance to photoinhibition in poplar trees. Plant Physiology, 109: 1047
Gupta A, Heinen J, Holaday A, Burke J, Allen R. 1993a.Increased resistance to oxidative stress in transgenic plants that overexpress chloroplastic Cu/Zn superoxide dismutase. Proceedings of the National Academy of Sciences, 90: 1629-1633
Gupta A, Webb R, Holaday A, Allen R (1993b) Overexpression of superoxide dismutase protects plants from oxidative stress (induction of ascorbate peroxidase in superoxide dismutase-overexpressing plants). Am Soc Plant Biol, pp 1067-1073
Henzler T, Steudle E. 2000.Transport and metabolic degradation of hydrogen peroxide in Chara corallina: model calculations and measurements with the pressure probe suggest transport of H2O2 across water channels. Journal of Experimental Botany, 51: 2053
Hodges D, Andrews C, Johnson D, Hamilton R. 1997.Antioxidant enzyme responses to chilling stress in differentially sensitive inbred maize lines. Journal of Experimental Botany, 48: 1105-1113
Joo J, Bae Y, Lee J. 2001.Role of auxin-induced reactive oxygen species in root gravitropism. Science's STKE, 126: 1055
Kim K, Kwon S, Lee H, Hur Y, Bang J, Kwak S. 2003.A novel oxidative stress-inducible peroxidase promoter from sweetpotato: molecular cloning and characterization in transgenic tobacco plants and cultured cells. Plant molecular biology, 51: 831-838
Kratsch H, Wise R. 2000.The ultrastructure of chilling stress. Plant, Cell & Environment, 23: 337-350
Kwon S, Jeong Y, Lee H, Kim J, Cho K, Allen R, Kwak S. 2002.Enhanced tolerances of transgenic tobacco plants expressing both superoxide dismutase and ascorbate peroxidase in chloroplasts against methyl viologen-mediated oxidative stress. Plant, Cell & Environment, 25: 873-882
Lee D, Lee C. 2000.Chilling stress-induced changes of antioxidant enzymes in the leaves of cucumber in gel enzyme activity assay. Plant Science, 159: 75-85
Lee S, Choi H, Suh S, Doo I, Oh K, Jeong Choi E, Schroeder Taylor A, Low P, Lee Y. 1999.Oligogalacturonic acid and chitosan reduce stomatal aperture by inducing the evolution of reactive oxygen species from guard cells of tomato and Commelina communis. Plant Physiology, 121: 147
Li S, Xue L, Xu S, Feng H, An L. 2007.Hydrogen peroxide involvement in formation and development of adventitious roots in cucumber. Plant Growth Regulation, 52: 173-180
Lim S, Kim Y, Kim S, Kwon S, Lee H, Kim J, Cho K, Paek K, Kwak S. 2007.Enhanced tolerance of transgenic sweetpotato plants that express both CuZnSOD and APX in chloroplasts to methyl viologen-mediated oxidative stress and chilling. Molecular Breeding, 19: 227-239
McInnis S, Desikan R, Hancock J, Hiscock S. 2006.Production of reactive oxygen species and reactive nitrogen species by angiosperm stigmas and pollen: potential signalling crosstalk? New Phytologist, 172: 221-228
McKersie B, Bowley S, Harjanto E, Leprince O. 1996.Water-deficit tolerance and field performance of transgenic alfalfa overexpressing superoxide dismutase. Plant Physiology, 111: 1177-1181
McKersie B, Chen Y, De Beus M, Bowley S, Bowler C, Inze D, D'halluin K, Botterman J. 1993.Superoxide dismutase enhances tolerance of freezing stress in transgenic alfalfa (Medicago sativa L.). Plant Physiology, 103: 1155
McKersie B, Murnaghan J, Jones K, Bowley S. 2000.Iron-superoxide dismutase expression in transgenic alfalfa increases winter survival without a detectable increase in photosynthetic oxidative stress tolerance. Plant Physiology, 122: 1427-1438
Mittler R. 2002.Oxidative stress, antioxidants and stress tolerance. Trends in Plant Science, 7: 405-410
Mittler R, Herr E, Orvar B, van Camp W, Willekens H, InzéD, Ellis B. 1999.Transgenic tobacco plants with reduced capability to detoxify reactive oxygen intermediates are hyperresponsive to pathogen infection. Proceedings of the National Academy of Sciences of the United States of America, 96: 14165
Mukherjee S, Choudhuri M. 1983.Determination of glycolate oxidase activity, H2O2 content and catalase activity. Physiol Plant, 58: 167-170
Neill S, Desikan R, Clarke A, Hurst R, Hancock J (2002a) Hydrogen peroxide and nitric oxide as signalling molecules in plants. Soc Experiment Biol, pp 1237-1247
Neill S, Desikan R, Hancock J. 2002b.Hydrogen peroxide signalling. Current Opinion in Plant Biology, 5: 388-395
Perl A, Perl-Treves R, Galili S, Aviv D, Shalgi E, Malkin S, Galun E. 1993.Enhanced oxidative-stress defense in transgenic potato expressing tomato Cu, Zn superoxide dismutases. TAG Theoretical and Applied Genetics, 85: 568-576
Pitcher L, Repetti P, Zilinskas B. 1994.Overproduction of ascorbate peroxidase protects transgenic tobacco against oxidative stress (abstract No. 623). Plant Physiol, 105: 116
Potikha T, Collins C, Johnson D, Delmer D, Levine A. 1999.The involvement of hydrogen peroxide in the differentiation of secondary walls in cotton fibers. Plant Physiology, 119: 849
Potocky M, Jones M, Bezvoda R, Smirnoff N,ársky V. 2007.Reactive oxygen species produced by NADPH oxidase are involved in pollen tube growth. Comparative Biochemistry and Physiology, Part A, 146: 269-270
Prasad T, Anderson M, Martin B, Stewart C. 1994.Evidence for chilling-induced oxidative stress in maize seedlings and a regulatory role for hydrogen peroxide. The Plant Cell Online, 6: 65-74
Ren D, Yang H, Zhang S. 2002.Cell death mediated by MAPK is associated with hydrogen peroxide production in Arabidopsis. Journal of Biological Chemistry, 277: 559-565
Shah K, Kumar R, Verma S, Dubey R.Effect of cadmium on lipid peroxidation, superoxide anion generation and activities of antioxidant enzymes in growing rice seedlings.Время, 22: 16
Slooten L, Capiau K, Van Camp W, Van Montagu M, Sybesma C, InzéD. 1995.Factors affecting the enhancement of oxidative stress tolerance in transgenic tobacco overexpressing manganese superoxide dismutase in the chloroplasts. Plant Physiology, 107: 737
Su G, Zhang W, Liu Y. 2006.Involvement of hydrogen peroxide generated by polyamine oxidative degradation in the development of lateral roots in soybean. Journal of Integrative Plant Biology, 48: 426
Svalheim, Robertsen B. 2006.Elicitation of H2O2 production in cucumber hypocotyl segments by oligo-1, 4-α-D-galacturonides and an oligo-β-glucan preparation from cell walls of Phythophthora megasperma f. sp. glycinea. Physiologia Plantarum, 88: 675-681
Svalheim O, Robertsen B. 1993.Elicitation of hydrogen peroxide production in cucumber hypocotyl segments by oligo-14-alpha-d-galacturonides and an oligo-beta-glucan preparation from cell walls of Phytophthora megasperma f. sp. glycinea. Physiol Plant, 88: 675-681
Tang L, Kwon S, Kim S, Kim J, Choi J, Cho K, Sung C, Kwak S, Lee H. 2006.Enhanced tolerance of transgenic potato plants expressing both superoxide dismutase and ascorbate peroxidase in chloroplasts against oxidative stress and high temperature. Plant Cell Reports, 25: 1380-1386
Tepperman J, Dunsmuir P. 1990.Transformed plants with elevated levels of chloroplastic SOD are not more resistant to superoxide toxicity. Plant molecular biology, 14: 501-511
Tiwari B, Belenghi B, Levine A. 2002.Oxidative stress increased respiration and generation of reactive oxygen species, resulting in ATP depletion, opening of mitochondrial permeability transition, and programmed cell death. Plant Physiology, 128: 1271
Van Camp W, Capiau K, Van Montagu M, Inze D, Slooten L (1996a) Enhancement of oxidative stress tolerance in transgenic tobacco plants overproducing Fe-superoxide dismutase in chloroplasts. Am Soc Plant Biol, pp 1703-1714
Van Camp W, Capiau K, Van Montagu M, Inze D, Slooten L. 1996b.Enhancement of oxidative stress tolerance in transgenic tobacco plants overproducing Fe-superoxide dismutase in chloroplasts. Plant Physiology, 112: 1703
Webb R, Allen R. 1996.Overexpression of pea cytosolic ascorbate peroxidase confers protection against oxidative stress in transgenic Nicotiana tabacum. Plant Physiol. Supplement, 111: 48
YU C, MURPHY T, SUNG W, LIN C. 2002.H2O2 treatment induces glutathione accumulation and chilling tolerance in mung bean. Functional plant biology, 29: 1081-1087
Zhang X, Zhang L, Dong F, Gao J, Galbraith D, Song C. 2001.Hydrogen peroxide is involved in abscisic acid-induced stomatal closure in Vicia faba. Plant Physiology, 126: 1438