莲子长寿机制的研究进展
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摘要
莲(Nelumbo nucifera Gaertn.)为莲科、莲属多年水生宿根草本植物。莲子生命力极强,在自然条件下的泥碳层中能存活千年,甚至在极端高温、高湿条件下仍能保持较高的萌发活力。随着莲基因组测序与基因注释的完善,莲子长寿机制的相关研究取得了一定进展。本文对莲子的结构形态、保护系统和基因组特性方面的研究进展进行了综述,阐明莲子的长寿机制,对后续研究方向进行了展望。
Nelumbo nucifera Gaertn. belongs to the aquatic perennial herbs of Nelumbonaceae.The N. nucifera seeds exhibit strong vitality and can survive for more than a thousand years under natural and even extreme conditions. With the improvement of N. nucifera genome sequencing and gene annotation,some progress has been made in the study of the longevity mechanism of N. nucifera seeds. In this paper,the longevity mechanism is summarized from aspects of structure, morphology, anti-aging, and self-repair, and the prospects and challenges of follow-up research are discussed.
Nelumbo nucifera Gaertn. belongs to the aquatic perennial herbs of Nelumbonaceae.The N. nucifera seeds exhibit strong vitality and can survive for more than a thousand years under natural and even extreme conditions. With the improvement of N. nucifera genome sequencing and gene annotation,some progress has been made in the study of the longevity mechanism of N. nucifera seeds. In this paper,the longevity mechanism is summarized from aspects of structure, morphology, anti-aging, and self-repair, and the prospects and challenges of follow-up research are discussed.
引文
[1]董新红,宋明.种子寿命研究进展[J].生物学杂志,2001,18(6):7-9.Dong XH,Song M. The progress of research on seed life[J]. Journal of Biology,2001,18(6):7-9.
[2]张行言,王其超.中国荷花品种图志[M].北京:中国林业出版社,2005:47-52.
[3] Ohga I. On the longevity of seeds of Nelumbo nucifera[J]. Plant J,1923,37:87-95.
[4] Shen-Miller J. Sacred lotus,the long-living fruits of China Antique[J]. Seed Sci Res,2002,12(3):131-143.
[5] Shen-Miller J,Turek J,Schopf JW,Tholandi M,Yang M,et al. Centuries-Old viable fruit of sacred lotus Nelumbo nucifera Gaertn var. China Antique[J]. Trop Plant Biol,2013,6(2-3):53-68.
[6] Buitink J,Leprince O. Intracellular glasses and seed survival in the dry state[J]. C R Biol,2008,331(10):788-795.
[7]周德龙,高建华,杨祎静,陈霭君,麦振龙.莲子壳营养成分分析及类黄酮抗氧化活性研究[J].安徽农业科学,2011,39(7):3968-3970.Zhou DL,Gao JH,Yang YJ,Chen AJ,Mai ZL. Analysis on the nutrient components of lotus seed shell and study on the antioxidation activity of flavonoid[J]. Journal of Anhui Agricultural Sciences,2011,39(7):3968-3970.
[8]黄迪惠,胡崇琳,Credy HM,谢笔钧,何慧,杨尔宁.莲子壳提取物中黄酮类物质的抗氧化活性及结构初探[J].食品科学,2009,30(23):209-213.Hang DH,Hu CL,Credy HM,Xie BJ,He H,Yang EN.Antioxidant acticity and structure of flavonoids from epicarp of Nelumbo nucifera Geartn.[J]. Food Science,2009,30(23):209-213.
[9] Mohamedyasseen Y, Barringer SA, Splittstoesser WE,Costanza S. The role of seed coats in seed viability[J].Bot Rev,1994,60(4):426-439
[10] Barros J,Serk H,Granlund I,Pesquet E. The cell biology of lignification in higherplants[J]. Ann Bot,2015,115(7):1053-1074.
[11]陈轩,周坚.莲子皮化学成分的初步分析[J].农业机械,2011(20):139-141.Chen X,Zhou J. Preliminary analysis of chemical constituents of lotus seed peel[J]. Farm Machinery,2011(10):139-141.
[12] Li JJ,Shi T,Huang LY,He DL,Yang PF,et al. Systematic transcriptomic analysis provides insights into lotus(Nelumbo Nucifera)seed development[J]. Plant Growth Regul,2018,86:339-350.
[13]曾绍校.莲子淀粉品质特性的研究与应用[D].福州:福建农林大学,2007.
[14]刘子凡.种子学[M].北京:化学工业出版社,2016:26-35.
[15] Leopold AC,Sun WQ,Bernallugo I. The glassy state in seeds:analysis and function[J]. Seed Sci Res,1994,4(3):267-274.
[16]左宝玉,唐崇钦,宋云,匡廷云,段续川.莲子胚为什么是绿色的?[J].植物杂志,1985(6):12.Zuo BY,Tang CQ,Song Y,Kuang TY,Duan XC. Why is lotus seed embryo green?[J]. Plants,1985(6):12.
[17]黄天芳,李长春,肖丽.对莲子胚芽变绿的探讨[J].中国园艺文摘,2011,27(11):16-17.Huang TF,Li CC,Xiao L. Dicussion on the greening of the lotus[J]. Chinese Horiculture Abstracts,2011,27(11):16-17.
[18] Ushimaru T,Hasegawa T,Amano T,Katayama M,Tanaka S,et al. Chloroplasts in seeds and dark-grown seedlings of lotus[J]. J Plant Physiol,2003,160(3):321-324.
[19]季宏伟,唐崇钦,李良璧,匡廷云.莲胚芽暗萌发过程中的光合系统发育研究[J].植物学报,2001,43(11):1129-1133.Ji HW,Tang CQ,Li LB,Kuang TY. Photosystem development in dark-grown lotus(Nelumbo nucifera)seedlings[J]. Bulletin of Botany,2001,43(11):1129-1133.
[20] Sano N,Rajjou L,North HM,Debeaujon I,Marionpoll A,et al. Staying alive:molecular aspects of seed longevity[J]. Plant Cell Physiol,2016,57(4):660-674.
[21] Verdier J,Buitink J. A regulatory network-based approach dissects late maturation processes related to the acquisition of desiccation tolerance and longevity of medicago truncatula seeds[J]. Plant Physiol,2013,163(2):757-774.
[22] Hyde EOC. The function of the hilum in some papilionaceae in relation to the ripening of the seed and the permeability of the testa[J]. Ann Bot,1954,18(2):241-256.
[23] Ganesh KJ,Song DP,Liu W,Han YY,Liu BL. Relationship between seed moisture content and acquisition of impermeability in Nelumbo nucifera(Nelumbonaceae)[J].Acta Bot Bras,2017,31(4):639-644.
[24] Groot SPC,Surki AA,De Vos RCH,Kodde J. Seed storage at elevated partial pressure of oxygen,a fast method for analysing seed ageing under dry conditions[J]. Ann Bot,2012,110(6):1149-1159.
[25] Bailly C. Active oxygen species and antioxidants in seed biology[J]. Seed Sci Res,2004,14(2):93-107.
[26] Veselovsky VA,Veselova TV. Lipid peroxidation,carbohydrate hydrolysis,and Amadori-Maillard reaction at early stages of dry seed aging[J]. Russ J Plant Physl,2012,59(6):811-817.
[27] Galland M,Huguet R,Arc E,Cueff G,Job D,et al. Dynamic proteomics emphasizes the importance of selective mRNA translation and protein turnover during Arabidopsis seed germination[J]. Mol Cell Proteomics,2014,13(1):252-268.
[28] Ray P,Huang BW,Tsuji Y. Reactive oxygen species(ROS)homeostasis and redox regulation in cellular signaling[J]. Cell Signal,2012,24(5):981-990.
[29] Kocsy G. Die or survive? Redox changes as seed viability markers[J]. Plant Cell Environ,2015,38(12):1008-1010.
[30] Ding YF,Cheng HY,Song SQ. Changes in extreme hightemperature tolerance and activities of antioxidant enzymes of sacred lotus seeds[J]. Sci China C Life Sci,2008,51(9):842-853.
[31] Chen HH,Chu P,Zhou YL,Ding Y,Huang SZ,et al. Ectopic expression of Nn PER1,a Nelumbo nucifera 1-cysteine peroxiredox in antioxidant,enhances seed longevity and stress tolerance in Arabidopsis[J]. Plant J,2016,88(4):608-619.
[32] Shen-Miller J,Lindner P,Xie Y,Villa S,Wooding K,et al.Thermal-stable proteins of fruit of long-living sacred lotus Nelumbo nucifera Gaertn var. China Antique.[J]. Trop Plant Biol,2013,6(2-3):69-84.
[33] Gill SS,Tuteja N. Reactive oxygen species and antioxidant machinery in abiotic stress tolerance in crop plants[J]. Plant Physiol Bioch,2010,48(12):909-930.
[34] Kleindt CK,Stracke R,Mehrtens F,Weisshaar B. Expression analysis of flavonoid biosynthesis genes during Arabidopsis thaliana silique and seed development with a primary focus on the proanthocyanidin biosynthetic pathway[J]. BMC Res Notes,2010,3(1):255.
[35] Yen GC,Duh PD,Su HJ. Antioxidant properties of lotus seed and its effect on DNA damage damage in human lymphocytes[J]. Food Chem,2005,89(3):379-385.
[36] Nguyen TP,Cueff G,Hegedus DD,Rajjou L,Bentsink L.A role for seed storage proteins in Arabidopsis seed longevity lymphocytes[J]. J Exp Bot,2015,66(20):6399-6413.
[37] Qi Y,Wang H,Zou Y,Liu C,Liu Y,et al. Over-expression of mitochondrial heat shock protein 70 suppresses programmed cell death in rice[J]. FEBS Lett,2011,585(1):231-239.
[38] Prietodapena P,Castao R,Almoguera C,Jordano J. Improved resistance to controlled deterioration in transgenic seeds[J]. Plant Physiol,2006,142(3):1102-1112.
[39] Banti V,Loreti E,Novi G,Santaniello A,Aipl A,et al.Heat acclimation and cross-tolerance against anoxia in Arabidopsis[J]. Plant Cell Environ,2010,31(7):1029-1037.
[40] Malik MK,Slovin JP,Hwang CH,Zimmerman JL. Modified expression of a carrot small heat shock protein gene,Hsp17. 7 results in increased or decreased thermotolerance[J]. Plant J,2010,20(1):89-99.
[41] Herald V. The impact of oxidative stress on Arabidopsis,mitochondria[J]. Plant J,2010,32(6):891-904.
[42] Ravanel S,Gakière B,Job D,Douce R. The specific features of methionine biosynthesis and metabolism in plants[J]. Proc Natl Acad Sci USA,1998,95(13):7805-7812.
[43] Schopf JW. The paleobiological record of photosynthesis[J]. Photosynth Res,2011,107(1):87-101.
[44] Shen-Miller J,Mudgett MB,Schopf JW,Clarke S,Berger R. Exceptional seed longevity and robust growth:ancient sacred lotus from China[J]. Am J Bot,1995,82(11):1367-1380.
[45] OgéL,Broyart C,Collet B,Godin B,Jallet D,et al. Protein damage and repair controlling seed vigor and longevity[J]. Methods Mol Biol,2017,773(773):369-384.
[46] Villa ST,Xu Q,Downie AB,Clarke SG. Arabidopsis protein repair L-isoaspartyl methyltransferases:predominant activities at lethal temperatures[J]. Physiol Plantarum,2010,128(4):581-592.
[47] Chu P,Chen H,Zhou Y,Li Y,Ding Y,et al. Proteomic and functional analyses of Nelumbo nucifera annexins involved in seed thermotolerance and germination vigor[J].Planta,2012,235(6):1271-1288.
[48] Dong C,Zheng X,Diao Y,Wang Y,Zhou M,et al. Molecular cloning and expression analysis of a catalase gene(Nn CAT)from Nelumbo nucifera[J]. Appl Biochm Biotechnol,2015,177(6):1216-1228.
[49] Cheng LB,Yang JJ,Yin L,Hui LC,Qian HM,et al. Transcription factor Nn DREB1 from lotus improved drought tolerance in transgenic Arabidopsis thaliana[J]. Biol Plantarum,2017,61(4):1-8.
[50] Wang Q,Guan Y,Wu Y,Chen H,Chen F,et al. Overexpression of a rice OsDREB gene increases salt,drought,and low temperature tolerance in both Arabidopsis and rice[J]. Plant Mol Biol,2008,67(6):589-602.
[51] Zhou Y,Chu P,Chen H,Li Y,Liu J,et al. Overexpression of Nelumbo nucifera metallothioneins and 3 enhances seed germination vigor in Arabidopsis[J]. Planta,2012,235(3):523-537.
[52] Rajjou L,Duval M,Gallardo K,Catusse J,Bally J,et al.Seed germination and vigor metallothionein[J]. Annu Rev Plant Biol,2012,63(3):507-533.
[53] Li YD. Study on the ABA content and SOD activity in ancient aotus and modern lotus seeds[J]. Chinese Bull Bot,2000,17(5):439-442.
[54] Kwak JM,Mori IC,Pei ZM,Leonhardt N,Torres MA,et al. NADPH oxidase AtrbohD and AtrbohF genes function in ROS dependent ABA signaling in Arabidopsis[J].Embo J,2014,22(22):2623-2633.
[55] Wang Y,Fan GY,Liu YM,Sun FM,Chi CC,et al. The sacred lotus genome provides insights into the evolution of flowering plants[J]. Plant J,2013,76(4):557-567.
[56] Nelson DR. Cytochrome P450 genes from the sacred lotus genome[J]. Trop Plant Biol,2013,6(2-3):138-151.
[2]张行言,王其超.中国荷花品种图志[M].北京:中国林业出版社,2005:47-52.
[3] Ohga I. On the longevity of seeds of Nelumbo nucifera[J]. Plant J,1923,37:87-95.
[4] Shen-Miller J. Sacred lotus,the long-living fruits of China Antique[J]. Seed Sci Res,2002,12(3):131-143.
[5] Shen-Miller J,Turek J,Schopf JW,Tholandi M,Yang M,et al. Centuries-Old viable fruit of sacred lotus Nelumbo nucifera Gaertn var. China Antique[J]. Trop Plant Biol,2013,6(2-3):53-68.
[6] Buitink J,Leprince O. Intracellular glasses and seed survival in the dry state[J]. C R Biol,2008,331(10):788-795.
[7]周德龙,高建华,杨祎静,陈霭君,麦振龙.莲子壳营养成分分析及类黄酮抗氧化活性研究[J].安徽农业科学,2011,39(7):3968-3970.Zhou DL,Gao JH,Yang YJ,Chen AJ,Mai ZL. Analysis on the nutrient components of lotus seed shell and study on the antioxidation activity of flavonoid[J]. Journal of Anhui Agricultural Sciences,2011,39(7):3968-3970.
[8]黄迪惠,胡崇琳,Credy HM,谢笔钧,何慧,杨尔宁.莲子壳提取物中黄酮类物质的抗氧化活性及结构初探[J].食品科学,2009,30(23):209-213.Hang DH,Hu CL,Credy HM,Xie BJ,He H,Yang EN.Antioxidant acticity and structure of flavonoids from epicarp of Nelumbo nucifera Geartn.[J]. Food Science,2009,30(23):209-213.
[9] Mohamedyasseen Y, Barringer SA, Splittstoesser WE,Costanza S. The role of seed coats in seed viability[J].Bot Rev,1994,60(4):426-439
[10] Barros J,Serk H,Granlund I,Pesquet E. The cell biology of lignification in higherplants[J]. Ann Bot,2015,115(7):1053-1074.
[11]陈轩,周坚.莲子皮化学成分的初步分析[J].农业机械,2011(20):139-141.Chen X,Zhou J. Preliminary analysis of chemical constituents of lotus seed peel[J]. Farm Machinery,2011(10):139-141.
[12] Li JJ,Shi T,Huang LY,He DL,Yang PF,et al. Systematic transcriptomic analysis provides insights into lotus(Nelumbo Nucifera)seed development[J]. Plant Growth Regul,2018,86:339-350.
[13]曾绍校.莲子淀粉品质特性的研究与应用[D].福州:福建农林大学,2007.
[14]刘子凡.种子学[M].北京:化学工业出版社,2016:26-35.
[15] Leopold AC,Sun WQ,Bernallugo I. The glassy state in seeds:analysis and function[J]. Seed Sci Res,1994,4(3):267-274.
[16]左宝玉,唐崇钦,宋云,匡廷云,段续川.莲子胚为什么是绿色的?[J].植物杂志,1985(6):12.Zuo BY,Tang CQ,Song Y,Kuang TY,Duan XC. Why is lotus seed embryo green?[J]. Plants,1985(6):12.
[17]黄天芳,李长春,肖丽.对莲子胚芽变绿的探讨[J].中国园艺文摘,2011,27(11):16-17.Huang TF,Li CC,Xiao L. Dicussion on the greening of the lotus[J]. Chinese Horiculture Abstracts,2011,27(11):16-17.
[18] Ushimaru T,Hasegawa T,Amano T,Katayama M,Tanaka S,et al. Chloroplasts in seeds and dark-grown seedlings of lotus[J]. J Plant Physiol,2003,160(3):321-324.
[19]季宏伟,唐崇钦,李良璧,匡廷云.莲胚芽暗萌发过程中的光合系统发育研究[J].植物学报,2001,43(11):1129-1133.Ji HW,Tang CQ,Li LB,Kuang TY. Photosystem development in dark-grown lotus(Nelumbo nucifera)seedlings[J]. Bulletin of Botany,2001,43(11):1129-1133.
[20] Sano N,Rajjou L,North HM,Debeaujon I,Marionpoll A,et al. Staying alive:molecular aspects of seed longevity[J]. Plant Cell Physiol,2016,57(4):660-674.
[21] Verdier J,Buitink J. A regulatory network-based approach dissects late maturation processes related to the acquisition of desiccation tolerance and longevity of medicago truncatula seeds[J]. Plant Physiol,2013,163(2):757-774.
[22] Hyde EOC. The function of the hilum in some papilionaceae in relation to the ripening of the seed and the permeability of the testa[J]. Ann Bot,1954,18(2):241-256.
[23] Ganesh KJ,Song DP,Liu W,Han YY,Liu BL. Relationship between seed moisture content and acquisition of impermeability in Nelumbo nucifera(Nelumbonaceae)[J].Acta Bot Bras,2017,31(4):639-644.
[24] Groot SPC,Surki AA,De Vos RCH,Kodde J. Seed storage at elevated partial pressure of oxygen,a fast method for analysing seed ageing under dry conditions[J]. Ann Bot,2012,110(6):1149-1159.
[25] Bailly C. Active oxygen species and antioxidants in seed biology[J]. Seed Sci Res,2004,14(2):93-107.
[26] Veselovsky VA,Veselova TV. Lipid peroxidation,carbohydrate hydrolysis,and Amadori-Maillard reaction at early stages of dry seed aging[J]. Russ J Plant Physl,2012,59(6):811-817.
[27] Galland M,Huguet R,Arc E,Cueff G,Job D,et al. Dynamic proteomics emphasizes the importance of selective mRNA translation and protein turnover during Arabidopsis seed germination[J]. Mol Cell Proteomics,2014,13(1):252-268.
[28] Ray P,Huang BW,Tsuji Y. Reactive oxygen species(ROS)homeostasis and redox regulation in cellular signaling[J]. Cell Signal,2012,24(5):981-990.
[29] Kocsy G. Die or survive? Redox changes as seed viability markers[J]. Plant Cell Environ,2015,38(12):1008-1010.
[30] Ding YF,Cheng HY,Song SQ. Changes in extreme hightemperature tolerance and activities of antioxidant enzymes of sacred lotus seeds[J]. Sci China C Life Sci,2008,51(9):842-853.
[31] Chen HH,Chu P,Zhou YL,Ding Y,Huang SZ,et al. Ectopic expression of Nn PER1,a Nelumbo nucifera 1-cysteine peroxiredox in antioxidant,enhances seed longevity and stress tolerance in Arabidopsis[J]. Plant J,2016,88(4):608-619.
[32] Shen-Miller J,Lindner P,Xie Y,Villa S,Wooding K,et al.Thermal-stable proteins of fruit of long-living sacred lotus Nelumbo nucifera Gaertn var. China Antique.[J]. Trop Plant Biol,2013,6(2-3):69-84.
[33] Gill SS,Tuteja N. Reactive oxygen species and antioxidant machinery in abiotic stress tolerance in crop plants[J]. Plant Physiol Bioch,2010,48(12):909-930.
[34] Kleindt CK,Stracke R,Mehrtens F,Weisshaar B. Expression analysis of flavonoid biosynthesis genes during Arabidopsis thaliana silique and seed development with a primary focus on the proanthocyanidin biosynthetic pathway[J]. BMC Res Notes,2010,3(1):255.
[35] Yen GC,Duh PD,Su HJ. Antioxidant properties of lotus seed and its effect on DNA damage damage in human lymphocytes[J]. Food Chem,2005,89(3):379-385.
[36] Nguyen TP,Cueff G,Hegedus DD,Rajjou L,Bentsink L.A role for seed storage proteins in Arabidopsis seed longevity lymphocytes[J]. J Exp Bot,2015,66(20):6399-6413.
[37] Qi Y,Wang H,Zou Y,Liu C,Liu Y,et al. Over-expression of mitochondrial heat shock protein 70 suppresses programmed cell death in rice[J]. FEBS Lett,2011,585(1):231-239.
[38] Prietodapena P,Castao R,Almoguera C,Jordano J. Improved resistance to controlled deterioration in transgenic seeds[J]. Plant Physiol,2006,142(3):1102-1112.
[39] Banti V,Loreti E,Novi G,Santaniello A,Aipl A,et al.Heat acclimation and cross-tolerance against anoxia in Arabidopsis[J]. Plant Cell Environ,2010,31(7):1029-1037.
[40] Malik MK,Slovin JP,Hwang CH,Zimmerman JL. Modified expression of a carrot small heat shock protein gene,Hsp17. 7 results in increased or decreased thermotolerance[J]. Plant J,2010,20(1):89-99.
[41] Herald V. The impact of oxidative stress on Arabidopsis,mitochondria[J]. Plant J,2010,32(6):891-904.
[42] Ravanel S,Gakière B,Job D,Douce R. The specific features of methionine biosynthesis and metabolism in plants[J]. Proc Natl Acad Sci USA,1998,95(13):7805-7812.
[43] Schopf JW. The paleobiological record of photosynthesis[J]. Photosynth Res,2011,107(1):87-101.
[44] Shen-Miller J,Mudgett MB,Schopf JW,Clarke S,Berger R. Exceptional seed longevity and robust growth:ancient sacred lotus from China[J]. Am J Bot,1995,82(11):1367-1380.
[45] OgéL,Broyart C,Collet B,Godin B,Jallet D,et al. Protein damage and repair controlling seed vigor and longevity[J]. Methods Mol Biol,2017,773(773):369-384.
[46] Villa ST,Xu Q,Downie AB,Clarke SG. Arabidopsis protein repair L-isoaspartyl methyltransferases:predominant activities at lethal temperatures[J]. Physiol Plantarum,2010,128(4):581-592.
[47] Chu P,Chen H,Zhou Y,Li Y,Ding Y,et al. Proteomic and functional analyses of Nelumbo nucifera annexins involved in seed thermotolerance and germination vigor[J].Planta,2012,235(6):1271-1288.
[48] Dong C,Zheng X,Diao Y,Wang Y,Zhou M,et al. Molecular cloning and expression analysis of a catalase gene(Nn CAT)from Nelumbo nucifera[J]. Appl Biochm Biotechnol,2015,177(6):1216-1228.
[49] Cheng LB,Yang JJ,Yin L,Hui LC,Qian HM,et al. Transcription factor Nn DREB1 from lotus improved drought tolerance in transgenic Arabidopsis thaliana[J]. Biol Plantarum,2017,61(4):1-8.
[50] Wang Q,Guan Y,Wu Y,Chen H,Chen F,et al. Overexpression of a rice OsDREB gene increases salt,drought,and low temperature tolerance in both Arabidopsis and rice[J]. Plant Mol Biol,2008,67(6):589-602.
[51] Zhou Y,Chu P,Chen H,Li Y,Liu J,et al. Overexpression of Nelumbo nucifera metallothioneins and 3 enhances seed germination vigor in Arabidopsis[J]. Planta,2012,235(3):523-537.
[52] Rajjou L,Duval M,Gallardo K,Catusse J,Bally J,et al.Seed germination and vigor metallothionein[J]. Annu Rev Plant Biol,2012,63(3):507-533.
[53] Li YD. Study on the ABA content and SOD activity in ancient aotus and modern lotus seeds[J]. Chinese Bull Bot,2000,17(5):439-442.
[54] Kwak JM,Mori IC,Pei ZM,Leonhardt N,Torres MA,et al. NADPH oxidase AtrbohD and AtrbohF genes function in ROS dependent ABA signaling in Arabidopsis[J].Embo J,2014,22(22):2623-2633.
[55] Wang Y,Fan GY,Liu YM,Sun FM,Chi CC,et al. The sacred lotus genome provides insights into the evolution of flowering plants[J]. Plant J,2013,76(4):557-567.
[56] Nelson DR. Cytochrome P450 genes from the sacred lotus genome[J]. Trop Plant Biol,2013,6(2-3):138-151.