光调控植物叶绿素生物合成的研究进展
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摘要
总结了光信号(光强、光质、光周期及昼夜节律中光暗变化)对叶绿素含量及其生物合成途径中相关基因的影响。重点归纳了光信号途径中主要转录因子对叶绿素合成基因的调控机制,强调了表观遗传修饰在光调控叶绿素合成中的重要作用,以期探索光调控叶绿素积累的有效途径及靶标分子,为利用基因工程和环境调控手段定向调控有重要生物学功能的叶绿素提供理论基础。
This review summarized the studies concerning with the effects of light signaling(such as light intensity, light quality, photoperiod and circadian rhythm) on Chl accumulation and its biosynthesis-related genes. The regulation mechanism of Chl biosynthesis-related genes by transcription factors in the light signaling pathway and histone epigenetic modifications were emphasized,which may help us to explore effective pathways and target molecules in light-regulation of Chl biosynthesis,and promote Chl accumulation via genetic engineering and environmental regulation.
This review summarized the studies concerning with the effects of light signaling(such as light intensity, light quality, photoperiod and circadian rhythm) on Chl accumulation and its biosynthesis-related genes. The regulation mechanism of Chl biosynthesis-related genes by transcription factors in the light signaling pathway and histone epigenetic modifications were emphasized,which may help us to explore effective pathways and target molecules in light-regulation of Chl biosynthesis,and promote Chl accumulation via genetic engineering and environmental regulation.
引文
Abbas N,Maurya J P,Senapati D,Gangappa S N,Chattopadhyay S.2014.Arabidopsis CAM7 and HY5 physically interact and directly bind to the HY5 promoter to regulate its expression and thereby promote photomorphogenesis.The Plant Cell,26(3):1036-1052.
Ali M B,Khandaker L,Oba S.2009.Comparative study on functional components,antioxidant activity and color parameters of selected colored leafy vegetables as affected by photoperiods.Journal of Food Agriculture and Environment,7(3):392-398.
Ampomah-Dwamena C,Thrimawithana A H,Dejnoprat S,Lewis D,Espley R V,Allan A C.2018.A kiwifruit(Actinidia deliciosa)R2R3-MYBtranscription factor modulates chlorophyll and carotenoid accumulation.New Phytologist,221(1):309-325.
Armstrong G A,Runge S,Frick G,Sperling U,Apel K.1995.Identification of NADPH:protochlorophyllide oxidoreductases A and B:a branched pathway for lightdependent chlorophyll biosynthesis in Arabidopsis thaliana.Plant Physiology,108(4):1505-1517.
Bach A,Krol A.2001.Effect of light quality on somatic embryogenesis in Hyacinthus orientalis L.Delft’s blue.Biological Bulletin of Poznan,38(1):103-107.
Banas A K,Labuz J,Sztatelman O,Gabrys H,Fiedor L.2011.Expression of enzymes involved in chlorophyll catabolism in Arabidopsis is light controlled.Plant Physiology,157(3):1497-1504.
Bao Shunshu,He Dongxian,Guo Shunxing.2007.Suitable photoperiod of dendrobium officinal in vitro under controlled environment.Journal of Agricultural Science and Technology,9(6):90-94.(in Chinese)鲍顺淑,贺冬仙,郭顺星.2007.可控环境下光照时间对铁皮石斛组培苗生长发育的影响.中国农业科技导报,9(6):90-94.
Barajas-López J de D,Kremnev D,Shaikhali J,Pi?as-Fernández A,Strand?.2013.PAPP5 is involved in the tetrapyrrole mediated plastid signalling during chloroplast development.PLoS ONE,8(3):e60305.
Barnes S A,Nishizawa N K,Quaggio R B,Whitelam G C,Chua N H.1996.Far-red light blocks greening of Arabidopsis seedlings via a phytochrome A-mediated change in plastid development.The Plant Cell,8(4):601-615.
Beale S I.2005.Green genes gleaned.Trends in Plant Science,10(7):309-312.
Benhamed M,Bertrand C,Servet C,Zhou D X.2006.Arabidopsis GCN5,HD1,and TAF1/HAF2 interact to regulate histone acetylation required for light-responsive gene expression.The Plant Cell,18(11):2893-2903.
Benhamed M,Martin-Magniette M L,Taconnat L,Bitton F,Servet C,De Clercq R,De Meyer B,Buysschaert C,Rom-bauts S,Villarroel R,Aubourg S,Beynon J,Bhalerao R P,Coupland G,Gruissem W,Menke F,Weisshaar B,Renou J P,Zhou D X,Hilson P.2008.Genome-scale Arabidopsis promoter array identifies targets of the histone acetyltransferase GCN5.The Plant Journal,56(3):493-504.
Binkert M,Kozma-Bognár L,Terecskei K,de Veylder L,Nagy F,Ulm R.2014.UV-B-responsive association of the Arabidopsis bZIP transcription factor ELONGATED HYPOCOTYL5 with target genes,including its own promoter.The Plant Cell,26(10):4200-4213.
Brand A,Borovsky Y,Hill T,Rahman K A,Bellalou A,van Deynze A,Paran I.2014.CaGLK2 regulates natural variation of chlorophyll content and fruit color in pepper fruit.Theoretical Applied Genetics,127(10):2139-2148.
Brouwer B,Gardestr?m P,Keech O.2014.In response to partial plant shading,the lack of phytochrome A does not directly induce leaf senescence but alters the fine-tuning of chlorophyll biosynthesis.Journal of Experimental Botany,65(14):4037-4049.
CataláR,Medina J,Salinas J.2011.Integration of low temperature and light signaling during cold acclimation response in Arabidopsis.Proceedings of the National Academy of Sciences of the United States of America,108(39):16475-16480.
Chakraborty N,Tripathy B C.1992.Involvement of singlet oxygen in 5-aminolevulinic acid-induced photodynamic damage of cucumber(Cucumis sativus L.)chloroplasts.Plant Physiology,98(1):7-11.
Charron J B,He H,Elling A A,Deng X W.2009.Dynamic landscapes of four histone modifications during deetiolation in Arabidopsis.The Plant Cell,21(12):3732-3748.
Chen Xiang-wei,Liu Shi-qi,Liu Qing,Cheng Bo,Feng Lei,Liu Jing-kai.2013.Effects of different light qualities on growth and photosynthetic characteristics of Pakchoi.Northern Horticulture,(22):1-4.(in Chinese)陈祥伟,刘世琦,刘庆,成波,冯磊,刘景凯.2013.不同LED光源对小白菜生长及光合特性的影响.北方园艺,(22):1-4.
Chi S H.2003.Effect of photoperiod shortening on the nutrient uptake and carbon metabolism of tomato and hot pepper seedlings grown hydroponically.Protected Horticulture and Plant Factory,12(3):121-126.
Dhanya Thomas T T,Puthur J T.2017.UV radiation priming:a means of amplifying the inherent potential for abiotic stress tolerance in crop plants.Environmental and Experimental Botany,138:57-66.
Dietzel L,Gla?er C,Liebers M,Hiekel S,Courtois F,Czarnecki O,Schlicke H,Zubo Y,Borner T,Mayer K,Grimm B,Pfannschmidt T.2015.Identification of early nuclear target genes of plastidial redox signals that trigger the long-term response of Arabidopsis to light quality shifts.Molecular Plant,8(8):1237-1252.
Dong Li-hua,Yang Yong,Han Qiao-hong,Jia Xue-jing,Yuan Ming.2015.Effect of circadian rhythm change on chlorophyll biosynthesis in Chlorophytum capense.Plant Science Journal,33(4):528-535.(in Chinese)董立花,杨勇,韩巧红,贾学静,袁明.2015.宽叶吊兰叶绿素生物合成的昼夜节律变化.植物科学学报,33(4):528-535.
Duanmu D,Casero D,Dent R M,Gallaher S,Yang W,Rockwell N C,Martin S S,Grossman A R,Lagarias J C.2013.Retrograde bilin signaling enables chlamydomonas greening and phototrophic survival.Proceedings of the National Academy of Science of the United States of America,110(9):3621-3626.
Du Hong-tao,Liu Shi-qi,Pu Gao-bin.2005.Effects of light qualities on growth and chlorophyll fluorescence parameters in leaves of color pepper seedling.Acta Agriculturae Boreali-occidentalis Sinica,14(1):41-45.(in Chinese)杜洪涛,刘世琦,蒲高斌.2005.光质对彩色甜椒幼苗生长及叶绿素荧光特性的影响.西北农业学报,14(1):41-45.
Espineda C E,Linford A S,Devine D,Brusslan J A.1999.The AtCAO gene,encoding chlorophyll a oxygenase,is required for chlorophyll b synthesis in Arabidopsis thaliana.Proceedings of the National Academy of Science of the United States of America,96(18):10507-10511.
Fan X X,Zang J,Xu Z G,Guo S R,Jiao X L,Liu X Y,Gao Y.2013.Effects of different light quality on growth,chlorophyll concentration and chlorophyll biosynthesis precursors of non-heading Chinese cabbage(Brassica campestris L.).Acta Physiologiae Plantarum 35(9):2721-2726.
Feng C M,Qiu Y,Van Buskirk E K,Yang E J,Chen M.2014.Light-regulated gene repositioning in Arabidopsis.Nature Communications,5:3027.
Fromme P,Melkozernov A,Jordan P,Krauss N.2003.Structure and function of photosystemⅠ:interaction with its soluble electron carriers and external antenna systems.FEBS Letters,555(1):40-44.
Fusada N,Masuda T,Kuroda H,Shiraishi T,Shimada H,Ohta H,Takamiya K.2000.NADPH-protochlorophyllide oxidoreductase in cucumber is encoded by a single gene and its expression is transcriptionally enhanced by illumination.Photosynthesis Research,64(2-3):147-154.
Golden S S,Ishiura M,Johnson C H,Kondo T.1997.Cyanobalcterial circadian rhythms.Annual Review of Plant Physiology and Plant Molecular Biology,48(1):327-354.
Guo L,Zhou J,Elling A A,Charron J B,Deng X W.2008.Histone modifications and expression of light regulated genes in Arabidopsis are cooperatively influenced by changing light conditions.Plant Physiology,147(4):2070-2083.
He Jun-jun,Yang Jing-ping,Yang Hu,Zhao Xing,Ye Xin-yi.2014.Effects of light intensity and nitrogen supply on the dynamic characteristics of leaf SPAD value of rice canopy.Journal of Zhejiang University(Agriculture and Life Sciences),40(5):495-504.(in Chinese)何俊俊,杨京平,杨虎,赵杏,叶欣怡.2014.光照及氮素水平对水稻冠层叶片SPAD值动态变化的影响.浙江大学学报(农业与生命科学版),40(5):495-504.
He Wei,Chen Dan-yan,Hu Xiao-ting,Wang Xiao-xu,Chen Le-han,Zhang Hai-chun,Yang Zhen-chao.2018.Effects of different photoperiods and photonflux ratios of red and blur LEDs on growth and development of tomato plants.Acta Agriculturae Boreali-occidentalis Sinica,27(4):562-570.(in Chinese)何蔚,陈丹艳,胡晓婷,王晓旭,陈乐涵,张海春,杨振超.2018.不同光周期与光质配比对番茄植株生长发育的影响.西北农业学报,27(4):562-570.
Hua Jin-song,Dai Hong-yan,Xia Ming-zhong.2009.Effects of different light Intensity on photosynthetic characteristic and yield characters of kidney bean.Acta Agriculturae Boreali-Occidentalis Sinica,18(2):136-140.(in Chinese)华劲松,戴红燕,夏明忠.2009.不同光照强度对芸豆光合特性及产量性状的影响.西北农业学报,18(2):136-140.
Huang L,Bonner B A,Castelfranco P A.1989.Regulation of 5-aminolevulinic acid(ALA)synthesis in developing chloroplast regulation of ALA-synthesizing capacity by phytochrome.Plant Physiology,90(3):1003-1008.
Huq E,Al-Sady B,Hudson M,Kim C,Apel K,Quail P H.2004.Phytochrome-interacting factor 1 is a critical bHLH regulator of chlorophyll biosynthesis.Science,305(5692):1937-1941.
IIag L L,Kumar A M,S?ll D.1994.Light regulation of chlorophyll biosynthesis at the level of 5-aminolevulinate formation in Arabidopsis.The Plant Cell,6(2):265-275.
Ishijima S,Uchibori A,Takagi H,Maki R,Ohnishi M.2003.Light-induced increase in free Mg2+concentration in spinach chloroplasts:measurement of free Mg2+by using a fluorescent probe and necessity of stromal alkalinization.Archives of Biochemistry&Biophysics,412(1):126-132.
Jang I C,Henriques R,Seo H S,Nagatani A,Chua N H.2010.Arabidopsis PHYTOCHROME INTERACTING FACTOR proteins promote phytochrome B polyubiquitination by COP1 E3 ligase in the nucleus.The Plant Cell,22(7):2370-2383.
Jiao Y,Lau O S,Deng X W.2007.Light-regulated transcriptional networks in higher plants.Nature Reviews Genetics,8(3):217-230.
Jing Yan-jun,Lin Rong-cheng.2017.Progress in plant light signal transduction in China.Chinese Bulletin of Botany,52(3):257-270.(in Chinese)景艳军,林荣呈.2017.我国植物光信号转导研究进展概述.植物学报,52(3):257-270.
Kindgren P,Noren L,Barajas-Lopez J D,Shaikhali J,Strand A.2012.Interplay between HEAT SHOCK PROTEIN 90 and HY5 controls PhANGexpression in response to the GUN5 plastid signal.Molecular Plant,5(4):901-913.
Kobayashi K,Amore T,Lazaro M.2013.Light-Emitting diodes(LEDs)for miniature hydroponic lettuce.Optics and Photonics Journal,3(1):74-77.
Koussevitzky S,Nott A,Mockler T C,Hong F,Sachettomartins G,Surpin M,Lim J,Mittler R,Chory J.2007.Signals from chloroplasts converge to regulate nuclear gene expression.Science,316(5825):715-719.
Kruse E,Grimm B,Beator J,Kloppstech K.1997.Developmental and circadian control of the capacity ford-aminolevulinic acid synthesis in green barley.Planta,202(2):235-241.
Lee S,Kim J H,Yoo E S,Lee C H,Hirochika H,An G.2005.Differential regulation of chlorophyll a oxygenase genes in rice.Plant Molecular Biology,57(6):805-818.
Lepisto A,Kangasjarvi S,Luomala E M,Brader G,Sipari N,Keranen M,Keinanen M,Rintamaki E.2009.Chloroplast NADPH-thioredoxin reductase interacts with photoperiodic development in Arabidopsis.Plant Physiology,149(3):1261-1276.
Li G,Siddiqui H,Teng Y,Lin R,Wan X Y,Li J,Lau O S,Ouyang X,Dai M,Wan J,Devlin P F,Deng X W,Wang H.2011.Coordinated transcriptional regulation underlying the circadian clock in Arabidopsis.Nature Cell Biology,13(5):616-622.
Li Han-sheng,Xu Yong.2014.Effect of light on synthesis of chlorophylls.Modern Agricultural Science and Technology,(21):161-164.(in Chinese)李汉生,徐永.2014.光照对叶绿素合成的影响.现代农业科技,(21):161-164.
Li H R,Li Y X,Deng H,Sun X C,Wang A Q,Tang X F,Gao Y F,Zhang N,Wang L H,Yang S Z,Liu Y S,Wang S H.2018.Tomato UV-Breceptor SlUVR8 mediates plant acclimation to UV-B radiation and enhances fruit chloroplast development via regulating SlGLK2.Scientific Reports,8(1):6097.
Li J,Terzaghi W,Deng X W.2012.Genomic basis for light control of plant development.Protein&Cell,3(2):106-116.
Li Y,Mukherjee I,Thum K E,Tanurdzic M,Katari M S,Obertello M,Edwards M B,McCombie W R,Martienssen R A,Coruzzi G M.2015.The histone methyltransferase SDG8 mediates the epigenetic modification of light and carbon responsive genes in plants.Genome Biology,16(1):79.
Lim J,Park J H,Jung S,Hwang D,Nam H G,Hong S.2018.Antagonistic roles of phyA and phyB in far-red light-dependent leaf senescence in Arabidopsis thaliana.Plant and Cell Physiology,59(9):1753-1764.
Liu Guo-shun,Qiao Xin-rong,Wang Fang,Yang Chao,Guo Qiao-yan,Yun Fei.2007.Effects of light intensity on photosynthetic capabilities,growth and quality of flue-cured tobacco.Acta Botanica Boreali-Occidentalia Sinica,27(9):1833-1837.(in Chinese)刘国顺,乔新荣,王芳,杨超,郭桥燕,云菲.2007.光照强度对烤烟光合特性及其生长和品质的影响.西北植物学报,27(9):1833-1837.
Liu Qing.2015.Effect of different photoperiod and light qualities on physiological characters and quality in strawberry[M.D.Dessertation].Tai’an:Shandong Agriculture University.(in Chinese)刘庆.2015.不同光周期及光质对草莓生理特性及品质的影响[硕士论文].泰安:山东农业大学,
Liu R,Xu Y H,Jiang S C,Lu K,Lu Y F,Feng X J,Wu Z,Liang S,Yu Y T,Wang X F,Zhang D P.2013a.Light-harvesting chlorophyll a/b-binding proteins,positively involved in abscisic acid signalling,require a transcription repressor,WRKY40,to balance their function.Journal of Experimental Botany,64(18):5443-5456.
Liu X,Chen C Y,Wang K C,Luo M,Tai R,Yuan L,Zhao M,Yang S,Tian G,Cui Y,Hsieh H L,Wu K.2013b.PHYTOCHROMEINTERACTING FACTOR3 associates with the histone deacetylase HDA15 in repression of chlorophyll biosynthesis and photosynthesis in etiolated Arabidopsis seedlings.The Plant Cell,25(4):1258-1273.
Liu Xiao-ying,Xu Zhi-gang,Chang Tao-tao,Guo Shi-rong.2010.Growth and photosynthesis of cherry tomato seedling exposed to different low light of LED light quality.Acta Botanica Boreali-Occidentalia Sinica,30(4):725-732.(in Chinese)刘晓英,徐志刚,常涛涛,郭世荣.2010.不同光质LED弱光对樱桃番茄植株形态和光合性能的影响.西北植物学报,30(4):725-732.
Lu Dong-yu,Cui Xin-tu,Huang Jing-rong,Zhao Fu-li.2006.Investigation on absorption spectrum of chlorophyll.College Physics,25(1):51-53.(in Chinese)卢东昱,崔新图,黄镜荣,赵福利.2006.叶绿素吸收光谱的观测.大学物理,25(1):51-53.
Mart?n G,Leivar P,Ludevid D,Tepperman J M,Quail P H,Monte E.2016.Phytochrome and retrograde signaling pathways converge to antagonistically regulate a light-induced transcriptional network.Nature Communications,(7):11431.
Ma Shao-ying,Li Sheng,Niu Jun-yi,Zhang Zhen,Liu Yuan,Xue Chong.2010.Effects of different LED light on physiological and biochemical characters of grape rootstock plantlets.Journal of Gansu Agricultural University,45(5):56-62.(in Chinese)马绍英,李胜,牛俊义,张真,刘媛,薛冲.2010.LED不同光质对葡萄砧木试管苗生理生化特性的影响.甘肃农业大学学报,45(5):56-62.
Masuda T,Fusada N,Oosawa N,Takamatsu K,Yamamoto Y Y,Ohto M,Nakamura K,Goto K,Shibata D,Shirano Y,Hayashi H,Kato T,Tabata S,Shimada H,Ohta H,Takamiya K.2003.Functional analysis of isoforms of NADPH:protochlorophyllide oxidoreductase(POR),PORB and PORC,in Arabidopsis thaliana.Plant&Cell Physiology,44(10):963-974.
Matsumoto F,Obayashi T,Sasaki-Sekimoto Y,Ohta H,Takamiya K,Masuda T.2004.Gene expression profiling of the tetrapyrrole metabolic pathway in Arabidopsis with a mini-array system.Plant Physiology,135(4):2379-2391.
McCormac A C,Terry M J.2004.The nuclear genes Lhcb and HEMA1 are differentially sensitive to plastid signals and suggest distinct roles for the GUN1 and GUN5 plastid-signalling pathways during de-etiolation.The Plant Journal,40(5):672-685.
Mock H P,Trainotti L,Kruse E,Grimm B.1995.Isolation,sequencing and expression of c DNA sequences encoding uroporphyrinogen decarboxylase from tobacco and barley.Plant Molecular Biology,28(2):245-256.
Moon J,Zhu L,Shen H,Huq E.2008.PIF1 directly and indirectly regulates chlorophyll biosynthesis to optimize the greening process in Arabidopsis.Proceedings of the National Academy of Sciences of the United States of America,105(27):9433-9438.
Ni Jiheng,Chen Xuehao,Chen Chunhong,Xu Qiang.2009.Effects of supplemental different light qualities on growth,photosynthesis,biomass partition and early yield of greenhouse cucumber.Scientia Agricultura Sinica,42(7):2615-2623.(in Chinese)倪纪恒,陈学好,陈春宏,徐强.2009.补充不同光质对温室黄瓜生长发育光合和期产量的影响.中国农业科学,42(7):2615-2623.
Ni W,Xu S L,Tepperman J M,Stanley D J,Maltby D A,Gross J D,Burlingame A L,Wang Z Y,Quail P H.2014.A mutually assured destruction mechanism attenuates light signaling in Arabidopsis.Science,344(6188):1160-1164.
Ni Z,Kim E D,Ha M,Lackey E,Liu J,Zhang Y,Sun Q,Chen Z J.2009.Altered circadian rhythms regulate growth vigour in hybrids and allopolyploids.Nature,457(7277):327-331.
Noren L,Kindgren P,Stachula P,Ruhl M,Eriksson M E,Hurry V,Strand A.2016.Circadian and plastid signaling pathways are integrated to ensure correct expression of the CBF and COR genes during photoperiodic growth.Plant Physiology,171(2):1392-1406.
Nyitrai P,Sarvari E,Keresztes A,Lang F.1994.Organization of thylakoid membranes in low light grown maize seedlings.Effect of lincomycin treatment.Journal of Plant Physiology,144(3):370-375.
Oda-Yamamizo C,Mitsuda N,Sakamoto S,Ogawa D,Ohme-Takagi M,Ohmiya A.2016.The NAC transcription factor ANAC046 is a positive regulator of chlorophyll degradation and senescence in Arabidopsis leaves.Scientific Reports,6:23609.
Osterlund M T,Hardtke C S,Wei N,Deng X W.2000.Targeted destabilization of HY5 during lightregulated development of Arabidopsis.Nature,405(6785):462-466.
Ouyang X,Li J,Li G,Li B,Chen B,Shen H,Huang X,Mo X,Wan X,Lin R,Li S,Wang H,Deng X W.2011.Genome-wide binding site analysis of FAR-RED ELONGATED HYPOCOTYL3 reveals its novel function in Arabidopsis development.The Plant Cell,23(7):2514-2535.
Papenbrock J,Mock H P,Kruse E,Grimm B.1999.Expression studies in tetrapyrrole biosynthesis:inverse maxima of magnesium chelatase and ferrochelatase activity during cyclic photoperiods.Planta,208(2):264-273.
Park J E,Jeong B R,Seungjae H.2013.Growth of lettuce in closed-type plant production system as affected by light intensity and photoperiod under influence of white LED light.Protected Horticulture and Plant Factory,22(3):228-233.
Pattanayak G K,Tripathy B C.2011.Overexpression of protochlorophyllide oxidoreductase C regulates oxidative stress in Arabidopsis.PLoS ONE,6(10):e26532.
Petroutsos D,Tokutsu R,Maruyama S,Flori S,Greiner A,Magneschi L.,Cusant L,Kottke T,Mittag M,Hegemann P,Finazzi G,Minagawa J.2016.A blue-light photoreceptor mediates the feedback regulation of photosynthesis.Nature,537(7621):563-566.
Powell A L,Nguyen C V,Hill T,Cheng K L,Figueroa-Balderas R,Aktas H,Ashrafi H,Pons C,Fernández-Mu?oz R,Vicente A,Lopez-Baltazar J,Barry C S,Liu Y,Chetelat R,Granell A,Van Deynze A,Giovannoni J J,Bennett A B.2012.Uniform ripening encodes a Golden 2-like transcription factor regulating tomato fruit chloroplast development.Science,336(6039):1711-1715.
Qiu K,Li Z,Yang Z P,Chen J Y,Wu S X,Zhu X Y,Gao S,Gao J,Ren G D,Kuai B,Zhou X.2015.EIN3 and ORE1 accelerate degreening during ethylene-mediated leaf senescence by directly activating chlorophyll catabolic genes in Arabidopsis.PLoS Genetics,11(7):e1005399.
Reid J D,Hunter C N.2004.Magnesium-dependent ATPase activity and cooperativity of magnesium chelatase from Synechocystis sp.PCC6803.Journal of Biological Chemistry,279(26):26893-26899.
Rosado D,Gramegna G,Cruz A,Lira B S,Freschi L,de Setta N,Rossi M.2016.Phytochrome interacting factors(PIFs)in Solanum lycopersicum:diversity,evolutionary history and expression profiling during different developmental processes.PLoS ONE,11(11):e0165929.
Rusaczonek A,Czarnocka W,Kacprzak S,WitońD,?lesak I,Szechyńska-Hebda M,Gawroński P,Karpiński S.2015.Role of phytochromes A and B in the regulation of cell death and acclimatory responses to UR stress in Arabidopsis thaliana.Journal of Experimental Botany,66(21):6679-6695.
Saebo A,Krekling T,Appelgren M.1995.Light quality affects photosynthesis and leaf anatomy of birch plantlets in vitro.Plant Cell Tissue&Organ Culture,41(2):177-185.
Sakuraba Y,Han S H,Lee S H,H?rtensteiner S,Paek N C.2016.Arabidopsis NAC016 promotes chlorophyll breakdown by directly upregulating STAYGREEN1 transcription.Plant Cell Reports,35(1):155-166.
Sakuraba Y,Jeong J,Kang M Y,Kim J,Paek N C,Choi G.2014.Phytochrome-interacting transcription factors PIF4 and PIF5 induce leaf senescence in Arabidopsis.Nature Communications,14(5):4636.
Sakuraba Y,Kim E Y,Han S H,Piao W,An G,Todaka D,Yamaguchi-Shinozaki K,Paek N C.2017.Rice Phytochrome-Interacting Factor-Like1(Os PIL1)is involved in the promotion of chlorophyll biosynthesis through feed-forward regulatory loops.Journal of Experimental Botany,68(15):4103-4114.
Schmid H C,Oster U,Kogel J,Lenz S,Rudiger W.2001.Cloning and characterisation of chlorophyll synthase from Avena sativa.Biological Chemistry,382(6):903-911.
Servet C,Conde e Silva N,Zhou D X.2010.Histone acetyl-transferase AtGCN5/HAG1 is a versatile regulator of developmental and inducible gene expression in Arabidopsis.Molecular Plant,3(4):670-677.
Shin J,Kim K,Kang H,Zulfugarov I S,Bae G,Lee C H,Lee D,Choi G.2009.Phytochromes promote seedling light responses by inhibiting four negatively-acting phytochrome-interacting factors.Proceedings of the National Academy of Sciences of the United States of America,106(18):7660-7665.
Shi Qing-hua,Zhu Zhu-jun,Ying Quan-shen,Qian Qiong-qiu.2005.Effects of excess Mn on photosynthesis characteristics in cucumber under different light intensity.Chinese Journal of Applied Ecology,16(6):1047-1050.(in Chinese)史庆华,朱祝军,应泉盛,钱琼秋.2005.不同光强下高锰对黄瓜光合作用特性的影响.应用生态学报,16(6):1047-1050.
Song Y,Yang C,Gao S,Zhang W,Li L,Kuai B.2014.Age-triggered and dark-induced leaf senescence require the bHLH transcription factors PIF3,4,and 5.Molecular Plant,7(12):1776-1787.
Stenbaek A,Jensen P E.2010.Redox regulation of chlorophyll biosynthesis.Phytochemistry,71(8-9):853-859.
Stephenson P G,Terry M J.2008.Light signalling pathways regulating the Mg-chelatase branchpoint of chlorophyll synthesis during de-etiolation in Arabidopsis thaliana.Photochemical&Photobiological Sciences,7(10):1243-1252.
Sui X L,Mao S L,Wang L H,Zhang B X,Zhang Z X.2012.Effect of low light on the characteristics of photosynthesis and chlorophyll a fluorescence during leaf development of sweet pepper.Journal of Integrative Agriculture,11(10):1633-1643.
Sun X,Feng P,Xu X,Guo H,Ma J,Chi W,Lin R,Lu C,Zhang L.2011.A chloroplast envelope-bound PHD transcription factor mediates chloroplast signals to the nucleus.Nature Communications,(2):477.
Tang W,Wang W,Chen D,Ji Q,Jing Y,Wang H,Lin R.2012.Transposase-derived proteins FHY3/FAR1 interact with PHYTOCHROME-INTERACTING FACTOR1 to regulate chlorophyll biosynthesis by modulating HEMB1 during deetiolation in Arabidopsis.The Plant Cell,24(5):1984-2000.
Thum K E,Kim M Y,Christopher D A,Mullet J E.2001.Cryptochrome 1,cryptochrome 2,and phytochrome A co-activate the chloroplast psbDblue light-responsive promoter.The Plant Cell,13(12):2747-2760.
Toledo-Ortiz G,Johansson H,Lee K P,Bou-Torrent J,Stewart K,Steel G,Rodríguez-Concepción M,Halliday K J.2014.The HY5-PIF regulatory module coordinates light and temperature control of photosynthetic gene transcription.PLoS Genetics,10(6):e1004416.
Tucker D J.1981.Phytochrome regulation of leaf senescence in cucumber and tomato.Plant Science Letters,23(1):103-108.
Tzvetkova-Chevolleau T,Franck F,Alawady A E,Dall'Osto L,Carrière F,Bassi R,Grimm B,Nussaume L,Havaux M.2007.The light stress-induced protein ELIP2 is a regulator of chlorophyll synthesis.The Plant Journal,50(5):795-809.
Ueda Minoru,Tanaka Ayumi,Sugimoto Kazuhiko,Shikanai Toshiharu,Nishimura Yoshiki.2014.ChlB requirement for chlorophyll biosynthesis under short photoperiod in Marchantia polymorpha L.Genome Biology Evolution,6(3):620-628.
Valdés A E,Rizzardi K,Johannesson H,Para A,Sund?s-Larsson A,Landberg K.2012.Arabidopsis thaliana TERMINAL FLOWER2 is involved in light-controlled signalling during seedling photomorphogenesis.Plant Cell and Environment,35(6):1013-1025.
Vothknecht U C,Kannangara C G,Von W D.1988.Barley glutamyl tRNA-Glu reductase:mutations affecting haem inhibition and enzyme activity.Phytochemistry,47(4):513-519.
Wang Z H,Hong X,Hu K K,Wang Y,Wang X X,Du S Y,Li Y,Hu D D,Cheng K,An B G,Li Y S.2017.Impaired magnesium protoporphyrin IX methyltransferase(ChlM)impedes chlorophyll synthesis and plant growth in rice.Frontiers in Plant Science,8:1-18.
Waters M T,Wang P,Korkaric M,Capper R G,Saunders N J,Langdale J A.2009.GLK transcription factors coordinate expression of the photosynthetic apparatus in Arabidopsis.The Plant Cell,21(4):1109-1128.
Xu G,Guo H,Zhang D,Chen D,Jiang Z,Lin R.2015.REVEILLE1 promotes NADPH:protochlorophyllide oxidoreductase A expression and seedling greening in Arabidopsis.Photosynthesis Research,126(2-3):331-340.
Xu Kai,Guo Yan-ping,Zang Shang-long.2005.Effect of light quality on photosynthesis and chlorophyll fluorescence in strawberry leaves.Scientia Agricultura Sinica,38(2):369-375.(in Chinese)徐凯,郭延平,张上隆.2005.不同光质对草莓叶片光合作用和叶绿素荧光的影响.中国农业科学,38(2):369-375.
Xu Li,Wei Hui,Qi Lian-dong,Hao Shu-qin,Liang Qing-ling,Yu Wen-yan,Liu Shi-qi.2010.Effects of different light qualities on growth and quality of Lactuca sativa L.China Fruit&Vegetable,(4):19-22.(in Chinese)许莉,尉辉,齐连东,郝树芹,梁庆玲,于文艳,刘世琦.2010.不同光质对叶用莴苣生长和品质的影响.中国果菜,(4):19-22.
Xu X,Chi W,Sun X,Feng P,Guo H,Li J,Lin R,Lu C,Wang H,Leister D,Zhang L.2016.Convergence of light and chloroplast signals for de-etiolation through ABI4-HY5 and COP1.Nature Plants,2(6):16066.
Xu X,Hotta C T,Dodd A N,Love J,Sharrock R,Lee Y W,Xie Q G,Johnson C H,Webb A A.2007.Distinct light and clock modulation of cytosolic free Ca2+oscillations and rhythmic CHLOROPHYLL A/B BINDING PROTEIN2 promoter activity in Arabidopsis.The Plant Cell,19(11):3474-3490.
Yang L,Lou Y,Peng Z,Zhao H,Sun H,Gao Z.2015.Molecular characterization and primary functional analysis of PeMPEC,a magnesium-protoporphyrin IX monomethyl ester cyclase gene of bamboo(Phyllostachys edulis).Plant Cell Reports,34(11):2001-2011.
Yang Xiao-jian,Liu Shi-qi,Zhang Zi-kun,Liu Zhong-liang,Ma Lin,Zhang Yu.2011.Effects of different light emitting diode sources on growth and chlorophyll fluorescence parameters in garlic seedling.China Vegetables,(6):62-67.(in Chinese)杨晓健,刘世琦,张自坤,刘中良,马琳,张宇.2011.不同LED光源对青蒜苗生长及叶绿素荧光特性的影响.中国蔬菜,(6):62-67.
Yin X R,Xie X L,Xia X J,Yu J Q,Ferguson I B,Giovannoni J J,Chen K S.2016.Involvement of an ethylene response factor in chlorophyll degradation during citrus fruit degreening.The Plant Journal,86(5):403-412.
Yuan M,Yuan S,Zhang Z W,Xu F,Chen Y E,Du J B,Lin H H.2010.Putative mutation mechanism and light responses of a protochlorophyllide oxidoreductase-less barley mutant NYB.Plant&Cell Physiology,51(8):1361-1371.
Yuan M,Zhang D W,Zhang Z W,Chen Y E,Yuan S,Guo Y R,Lin H H.2012.Assembly of NADPH:protochlorophyllide oxidoreductase complex is needed for effective greening of barley seedlings.Journal of Plant Physiology,169(13):1311-1316.
Yuan S,Zhang Z W,Zheng C,Zhao Z Y,Wang Y,Feng L Y,Niu G,Wang C Q,Wang J H,Feng H,Xu F,Bao F,Hu Y,Cao Y,Ma L,Wang H,Kong D D,Xiao W,Lin H H,He Y.2016.Arabidopsis cryptochrome 1 functions in nitrogen regulation of flowering.Proceedings of the National Academy of Sciences of the United States of America,113(27):7661-7666.
Yuan Y J,Mei L H,Wu M B,Wei W,Shan W,Gong Z H,Zhang Q,Yang F Q,Yan F,Zhang Q,Luo Y Q,Xu X,Zhang W F,Miao MJ,Lu W J,Li Z G,Deng W.2018.SlARF10,an auxin response factor,is involved in chlorophyll and sugar accumulation during tomato fruit development.Journal of Experimental Botany,69(22):5507-5518.
Zha Lingyan,Liu Wenke.2017.Effects of light quality and photoperiod on growth and yield of cherry radish grown under red plus blue LEDs.China Illuminating Engineering Journal,28(6):83-91.(in Chinese)查凌雁,刘文科.2017.LED红蓝光质比及其光周期对樱桃萝卜品种生长与产量的影响.照明工程学报,28(6):83-91.
Zhang D,Li Y,Zhang X,Zha P,Lin R.2017.The SWI2/SNF2 chromatin-remodeling ATPase BRAHMA regulates chlorophyll biosynthesis in Arabidopsis.Molecular Plant,10(1):155-167.
Zhang D W,Yuan S,Xu F,Zhu F,Yuan M,Ye H X,Guo H Q,Lv X,Yin Y,Lin H H.2016.Light intensity affects chlorophyll synthesis during greening process by metabolite signal from mitochondrial alternative oxidase in Arabidopsis.Plant Cell&Environment,39(1):12-25.
Zhang Y,Liu Z,Chen Y,He J X,Bi Y.2015.PHYTOCHROME-INTERACTING FACTOR 5(PIF5)positively regulates dark-induced senescence and chlorophyll degradation in Arabidopsis.Plant Science,237:57-68.
Zhang Z W,Feng L Y,Cheng J,Tang H,Xu F,Zhu F,Zhao Z Y,Yuan M,Chen Y E,Wang J H,Yuan S,Lin H H.2013.The roles of two transcription factors,ABI4 and CBFA,in ABA and plastid signalling and stress responses.Plant Molecular Biology,83(4-5):445-458.
Zhao J,Zhou J J,Wang Y Y,Gu J W,Xie X Z.2013.Positive regulation of phytochrome B on chlorophyll biosynthesis and chloroplast development in rice.Rice Science,20(4):243-248.
Zhong S,Zhao M,Shi T,Shi H,An F,Zhao Q,Guo H.2009.EIN3/EIL1 cooperate with PIF1 to prevent photooxidation and to promote greening of Arabidopsis seedlings.Proceedings of the National Academy of Sciences of the United States of America,106(50):21431-21436.
Zhu X,Chen J,Xie Z,Gao J,Ren G,Gao S,Zhou X,Kuai B.2015.Jasmonic acid promotes degreening via MYC2/3/4-and ANAC019/055/072-mediated regulation of major chlorophyll catabolic genes.The Plant Journal,84(3):597-610.
Ali M B,Khandaker L,Oba S.2009.Comparative study on functional components,antioxidant activity and color parameters of selected colored leafy vegetables as affected by photoperiods.Journal of Food Agriculture and Environment,7(3):392-398.
Ampomah-Dwamena C,Thrimawithana A H,Dejnoprat S,Lewis D,Espley R V,Allan A C.2018.A kiwifruit(Actinidia deliciosa)R2R3-MYBtranscription factor modulates chlorophyll and carotenoid accumulation.New Phytologist,221(1):309-325.
Armstrong G A,Runge S,Frick G,Sperling U,Apel K.1995.Identification of NADPH:protochlorophyllide oxidoreductases A and B:a branched pathway for lightdependent chlorophyll biosynthesis in Arabidopsis thaliana.Plant Physiology,108(4):1505-1517.
Bach A,Krol A.2001.Effect of light quality on somatic embryogenesis in Hyacinthus orientalis L.Delft’s blue.Biological Bulletin of Poznan,38(1):103-107.
Banas A K,Labuz J,Sztatelman O,Gabrys H,Fiedor L.2011.Expression of enzymes involved in chlorophyll catabolism in Arabidopsis is light controlled.Plant Physiology,157(3):1497-1504.
Bao Shunshu,He Dongxian,Guo Shunxing.2007.Suitable photoperiod of dendrobium officinal in vitro under controlled environment.Journal of Agricultural Science and Technology,9(6):90-94.(in Chinese)鲍顺淑,贺冬仙,郭顺星.2007.可控环境下光照时间对铁皮石斛组培苗生长发育的影响.中国农业科技导报,9(6):90-94.
Barajas-López J de D,Kremnev D,Shaikhali J,Pi?as-Fernández A,Strand?.2013.PAPP5 is involved in the tetrapyrrole mediated plastid signalling during chloroplast development.PLoS ONE,8(3):e60305.
Barnes S A,Nishizawa N K,Quaggio R B,Whitelam G C,Chua N H.1996.Far-red light blocks greening of Arabidopsis seedlings via a phytochrome A-mediated change in plastid development.The Plant Cell,8(4):601-615.
Beale S I.2005.Green genes gleaned.Trends in Plant Science,10(7):309-312.
Benhamed M,Bertrand C,Servet C,Zhou D X.2006.Arabidopsis GCN5,HD1,and TAF1/HAF2 interact to regulate histone acetylation required for light-responsive gene expression.The Plant Cell,18(11):2893-2903.
Benhamed M,Martin-Magniette M L,Taconnat L,Bitton F,Servet C,De Clercq R,De Meyer B,Buysschaert C,Rom-bauts S,Villarroel R,Aubourg S,Beynon J,Bhalerao R P,Coupland G,Gruissem W,Menke F,Weisshaar B,Renou J P,Zhou D X,Hilson P.2008.Genome-scale Arabidopsis promoter array identifies targets of the histone acetyltransferase GCN5.The Plant Journal,56(3):493-504.
Binkert M,Kozma-Bognár L,Terecskei K,de Veylder L,Nagy F,Ulm R.2014.UV-B-responsive association of the Arabidopsis bZIP transcription factor ELONGATED HYPOCOTYL5 with target genes,including its own promoter.The Plant Cell,26(10):4200-4213.
Brand A,Borovsky Y,Hill T,Rahman K A,Bellalou A,van Deynze A,Paran I.2014.CaGLK2 regulates natural variation of chlorophyll content and fruit color in pepper fruit.Theoretical Applied Genetics,127(10):2139-2148.
Brouwer B,Gardestr?m P,Keech O.2014.In response to partial plant shading,the lack of phytochrome A does not directly induce leaf senescence but alters the fine-tuning of chlorophyll biosynthesis.Journal of Experimental Botany,65(14):4037-4049.
CataláR,Medina J,Salinas J.2011.Integration of low temperature and light signaling during cold acclimation response in Arabidopsis.Proceedings of the National Academy of Sciences of the United States of America,108(39):16475-16480.
Chakraborty N,Tripathy B C.1992.Involvement of singlet oxygen in 5-aminolevulinic acid-induced photodynamic damage of cucumber(Cucumis sativus L.)chloroplasts.Plant Physiology,98(1):7-11.
Charron J B,He H,Elling A A,Deng X W.2009.Dynamic landscapes of four histone modifications during deetiolation in Arabidopsis.The Plant Cell,21(12):3732-3748.
Chen Xiang-wei,Liu Shi-qi,Liu Qing,Cheng Bo,Feng Lei,Liu Jing-kai.2013.Effects of different light qualities on growth and photosynthetic characteristics of Pakchoi.Northern Horticulture,(22):1-4.(in Chinese)陈祥伟,刘世琦,刘庆,成波,冯磊,刘景凯.2013.不同LED光源对小白菜生长及光合特性的影响.北方园艺,(22):1-4.
Chi S H.2003.Effect of photoperiod shortening on the nutrient uptake and carbon metabolism of tomato and hot pepper seedlings grown hydroponically.Protected Horticulture and Plant Factory,12(3):121-126.
Dhanya Thomas T T,Puthur J T.2017.UV radiation priming:a means of amplifying the inherent potential for abiotic stress tolerance in crop plants.Environmental and Experimental Botany,138:57-66.
Dietzel L,Gla?er C,Liebers M,Hiekel S,Courtois F,Czarnecki O,Schlicke H,Zubo Y,Borner T,Mayer K,Grimm B,Pfannschmidt T.2015.Identification of early nuclear target genes of plastidial redox signals that trigger the long-term response of Arabidopsis to light quality shifts.Molecular Plant,8(8):1237-1252.
Dong Li-hua,Yang Yong,Han Qiao-hong,Jia Xue-jing,Yuan Ming.2015.Effect of circadian rhythm change on chlorophyll biosynthesis in Chlorophytum capense.Plant Science Journal,33(4):528-535.(in Chinese)董立花,杨勇,韩巧红,贾学静,袁明.2015.宽叶吊兰叶绿素生物合成的昼夜节律变化.植物科学学报,33(4):528-535.
Duanmu D,Casero D,Dent R M,Gallaher S,Yang W,Rockwell N C,Martin S S,Grossman A R,Lagarias J C.2013.Retrograde bilin signaling enables chlamydomonas greening and phototrophic survival.Proceedings of the National Academy of Science of the United States of America,110(9):3621-3626.
Du Hong-tao,Liu Shi-qi,Pu Gao-bin.2005.Effects of light qualities on growth and chlorophyll fluorescence parameters in leaves of color pepper seedling.Acta Agriculturae Boreali-occidentalis Sinica,14(1):41-45.(in Chinese)杜洪涛,刘世琦,蒲高斌.2005.光质对彩色甜椒幼苗生长及叶绿素荧光特性的影响.西北农业学报,14(1):41-45.
Espineda C E,Linford A S,Devine D,Brusslan J A.1999.The AtCAO gene,encoding chlorophyll a oxygenase,is required for chlorophyll b synthesis in Arabidopsis thaliana.Proceedings of the National Academy of Science of the United States of America,96(18):10507-10511.
Fan X X,Zang J,Xu Z G,Guo S R,Jiao X L,Liu X Y,Gao Y.2013.Effects of different light quality on growth,chlorophyll concentration and chlorophyll biosynthesis precursors of non-heading Chinese cabbage(Brassica campestris L.).Acta Physiologiae Plantarum 35(9):2721-2726.
Feng C M,Qiu Y,Van Buskirk E K,Yang E J,Chen M.2014.Light-regulated gene repositioning in Arabidopsis.Nature Communications,5:3027.
Fromme P,Melkozernov A,Jordan P,Krauss N.2003.Structure and function of photosystemⅠ:interaction with its soluble electron carriers and external antenna systems.FEBS Letters,555(1):40-44.
Fusada N,Masuda T,Kuroda H,Shiraishi T,Shimada H,Ohta H,Takamiya K.2000.NADPH-protochlorophyllide oxidoreductase in cucumber is encoded by a single gene and its expression is transcriptionally enhanced by illumination.Photosynthesis Research,64(2-3):147-154.
Golden S S,Ishiura M,Johnson C H,Kondo T.1997.Cyanobalcterial circadian rhythms.Annual Review of Plant Physiology and Plant Molecular Biology,48(1):327-354.
Guo L,Zhou J,Elling A A,Charron J B,Deng X W.2008.Histone modifications and expression of light regulated genes in Arabidopsis are cooperatively influenced by changing light conditions.Plant Physiology,147(4):2070-2083.
He Jun-jun,Yang Jing-ping,Yang Hu,Zhao Xing,Ye Xin-yi.2014.Effects of light intensity and nitrogen supply on the dynamic characteristics of leaf SPAD value of rice canopy.Journal of Zhejiang University(Agriculture and Life Sciences),40(5):495-504.(in Chinese)何俊俊,杨京平,杨虎,赵杏,叶欣怡.2014.光照及氮素水平对水稻冠层叶片SPAD值动态变化的影响.浙江大学学报(农业与生命科学版),40(5):495-504.
He Wei,Chen Dan-yan,Hu Xiao-ting,Wang Xiao-xu,Chen Le-han,Zhang Hai-chun,Yang Zhen-chao.2018.Effects of different photoperiods and photonflux ratios of red and blur LEDs on growth and development of tomato plants.Acta Agriculturae Boreali-occidentalis Sinica,27(4):562-570.(in Chinese)何蔚,陈丹艳,胡晓婷,王晓旭,陈乐涵,张海春,杨振超.2018.不同光周期与光质配比对番茄植株生长发育的影响.西北农业学报,27(4):562-570.
Hua Jin-song,Dai Hong-yan,Xia Ming-zhong.2009.Effects of different light Intensity on photosynthetic characteristic and yield characters of kidney bean.Acta Agriculturae Boreali-Occidentalis Sinica,18(2):136-140.(in Chinese)华劲松,戴红燕,夏明忠.2009.不同光照强度对芸豆光合特性及产量性状的影响.西北农业学报,18(2):136-140.
Huang L,Bonner B A,Castelfranco P A.1989.Regulation of 5-aminolevulinic acid(ALA)synthesis in developing chloroplast regulation of ALA-synthesizing capacity by phytochrome.Plant Physiology,90(3):1003-1008.
Huq E,Al-Sady B,Hudson M,Kim C,Apel K,Quail P H.2004.Phytochrome-interacting factor 1 is a critical bHLH regulator of chlorophyll biosynthesis.Science,305(5692):1937-1941.
IIag L L,Kumar A M,S?ll D.1994.Light regulation of chlorophyll biosynthesis at the level of 5-aminolevulinate formation in Arabidopsis.The Plant Cell,6(2):265-275.
Ishijima S,Uchibori A,Takagi H,Maki R,Ohnishi M.2003.Light-induced increase in free Mg2+concentration in spinach chloroplasts:measurement of free Mg2+by using a fluorescent probe and necessity of stromal alkalinization.Archives of Biochemistry&Biophysics,412(1):126-132.
Jang I C,Henriques R,Seo H S,Nagatani A,Chua N H.2010.Arabidopsis PHYTOCHROME INTERACTING FACTOR proteins promote phytochrome B polyubiquitination by COP1 E3 ligase in the nucleus.The Plant Cell,22(7):2370-2383.
Jiao Y,Lau O S,Deng X W.2007.Light-regulated transcriptional networks in higher plants.Nature Reviews Genetics,8(3):217-230.
Jing Yan-jun,Lin Rong-cheng.2017.Progress in plant light signal transduction in China.Chinese Bulletin of Botany,52(3):257-270.(in Chinese)景艳军,林荣呈.2017.我国植物光信号转导研究进展概述.植物学报,52(3):257-270.
Kindgren P,Noren L,Barajas-Lopez J D,Shaikhali J,Strand A.2012.Interplay between HEAT SHOCK PROTEIN 90 and HY5 controls PhANGexpression in response to the GUN5 plastid signal.Molecular Plant,5(4):901-913.
Kobayashi K,Amore T,Lazaro M.2013.Light-Emitting diodes(LEDs)for miniature hydroponic lettuce.Optics and Photonics Journal,3(1):74-77.
Koussevitzky S,Nott A,Mockler T C,Hong F,Sachettomartins G,Surpin M,Lim J,Mittler R,Chory J.2007.Signals from chloroplasts converge to regulate nuclear gene expression.Science,316(5825):715-719.
Kruse E,Grimm B,Beator J,Kloppstech K.1997.Developmental and circadian control of the capacity ford-aminolevulinic acid synthesis in green barley.Planta,202(2):235-241.
Lee S,Kim J H,Yoo E S,Lee C H,Hirochika H,An G.2005.Differential regulation of chlorophyll a oxygenase genes in rice.Plant Molecular Biology,57(6):805-818.
Lepisto A,Kangasjarvi S,Luomala E M,Brader G,Sipari N,Keranen M,Keinanen M,Rintamaki E.2009.Chloroplast NADPH-thioredoxin reductase interacts with photoperiodic development in Arabidopsis.Plant Physiology,149(3):1261-1276.
Li G,Siddiqui H,Teng Y,Lin R,Wan X Y,Li J,Lau O S,Ouyang X,Dai M,Wan J,Devlin P F,Deng X W,Wang H.2011.Coordinated transcriptional regulation underlying the circadian clock in Arabidopsis.Nature Cell Biology,13(5):616-622.
Li Han-sheng,Xu Yong.2014.Effect of light on synthesis of chlorophylls.Modern Agricultural Science and Technology,(21):161-164.(in Chinese)李汉生,徐永.2014.光照对叶绿素合成的影响.现代农业科技,(21):161-164.
Li H R,Li Y X,Deng H,Sun X C,Wang A Q,Tang X F,Gao Y F,Zhang N,Wang L H,Yang S Z,Liu Y S,Wang S H.2018.Tomato UV-Breceptor SlUVR8 mediates plant acclimation to UV-B radiation and enhances fruit chloroplast development via regulating SlGLK2.Scientific Reports,8(1):6097.
Li J,Terzaghi W,Deng X W.2012.Genomic basis for light control of plant development.Protein&Cell,3(2):106-116.
Li Y,Mukherjee I,Thum K E,Tanurdzic M,Katari M S,Obertello M,Edwards M B,McCombie W R,Martienssen R A,Coruzzi G M.2015.The histone methyltransferase SDG8 mediates the epigenetic modification of light and carbon responsive genes in plants.Genome Biology,16(1):79.
Lim J,Park J H,Jung S,Hwang D,Nam H G,Hong S.2018.Antagonistic roles of phyA and phyB in far-red light-dependent leaf senescence in Arabidopsis thaliana.Plant and Cell Physiology,59(9):1753-1764.
Liu Guo-shun,Qiao Xin-rong,Wang Fang,Yang Chao,Guo Qiao-yan,Yun Fei.2007.Effects of light intensity on photosynthetic capabilities,growth and quality of flue-cured tobacco.Acta Botanica Boreali-Occidentalia Sinica,27(9):1833-1837.(in Chinese)刘国顺,乔新荣,王芳,杨超,郭桥燕,云菲.2007.光照强度对烤烟光合特性及其生长和品质的影响.西北植物学报,27(9):1833-1837.
Liu Qing.2015.Effect of different photoperiod and light qualities on physiological characters and quality in strawberry[M.D.Dessertation].Tai’an:Shandong Agriculture University.(in Chinese)刘庆.2015.不同光周期及光质对草莓生理特性及品质的影响[硕士论文].泰安:山东农业大学,
Liu R,Xu Y H,Jiang S C,Lu K,Lu Y F,Feng X J,Wu Z,Liang S,Yu Y T,Wang X F,Zhang D P.2013a.Light-harvesting chlorophyll a/b-binding proteins,positively involved in abscisic acid signalling,require a transcription repressor,WRKY40,to balance their function.Journal of Experimental Botany,64(18):5443-5456.
Liu X,Chen C Y,Wang K C,Luo M,Tai R,Yuan L,Zhao M,Yang S,Tian G,Cui Y,Hsieh H L,Wu K.2013b.PHYTOCHROMEINTERACTING FACTOR3 associates with the histone deacetylase HDA15 in repression of chlorophyll biosynthesis and photosynthesis in etiolated Arabidopsis seedlings.The Plant Cell,25(4):1258-1273.
Liu Xiao-ying,Xu Zhi-gang,Chang Tao-tao,Guo Shi-rong.2010.Growth and photosynthesis of cherry tomato seedling exposed to different low light of LED light quality.Acta Botanica Boreali-Occidentalia Sinica,30(4):725-732.(in Chinese)刘晓英,徐志刚,常涛涛,郭世荣.2010.不同光质LED弱光对樱桃番茄植株形态和光合性能的影响.西北植物学报,30(4):725-732.
Lu Dong-yu,Cui Xin-tu,Huang Jing-rong,Zhao Fu-li.2006.Investigation on absorption spectrum of chlorophyll.College Physics,25(1):51-53.(in Chinese)卢东昱,崔新图,黄镜荣,赵福利.2006.叶绿素吸收光谱的观测.大学物理,25(1):51-53.
Mart?n G,Leivar P,Ludevid D,Tepperman J M,Quail P H,Monte E.2016.Phytochrome and retrograde signaling pathways converge to antagonistically regulate a light-induced transcriptional network.Nature Communications,(7):11431.
Ma Shao-ying,Li Sheng,Niu Jun-yi,Zhang Zhen,Liu Yuan,Xue Chong.2010.Effects of different LED light on physiological and biochemical characters of grape rootstock plantlets.Journal of Gansu Agricultural University,45(5):56-62.(in Chinese)马绍英,李胜,牛俊义,张真,刘媛,薛冲.2010.LED不同光质对葡萄砧木试管苗生理生化特性的影响.甘肃农业大学学报,45(5):56-62.
Masuda T,Fusada N,Oosawa N,Takamatsu K,Yamamoto Y Y,Ohto M,Nakamura K,Goto K,Shibata D,Shirano Y,Hayashi H,Kato T,Tabata S,Shimada H,Ohta H,Takamiya K.2003.Functional analysis of isoforms of NADPH:protochlorophyllide oxidoreductase(POR),PORB and PORC,in Arabidopsis thaliana.Plant&Cell Physiology,44(10):963-974.
Matsumoto F,Obayashi T,Sasaki-Sekimoto Y,Ohta H,Takamiya K,Masuda T.2004.Gene expression profiling of the tetrapyrrole metabolic pathway in Arabidopsis with a mini-array system.Plant Physiology,135(4):2379-2391.
McCormac A C,Terry M J.2004.The nuclear genes Lhcb and HEMA1 are differentially sensitive to plastid signals and suggest distinct roles for the GUN1 and GUN5 plastid-signalling pathways during de-etiolation.The Plant Journal,40(5):672-685.
Mock H P,Trainotti L,Kruse E,Grimm B.1995.Isolation,sequencing and expression of c DNA sequences encoding uroporphyrinogen decarboxylase from tobacco and barley.Plant Molecular Biology,28(2):245-256.
Moon J,Zhu L,Shen H,Huq E.2008.PIF1 directly and indirectly regulates chlorophyll biosynthesis to optimize the greening process in Arabidopsis.Proceedings of the National Academy of Sciences of the United States of America,105(27):9433-9438.
Ni Jiheng,Chen Xuehao,Chen Chunhong,Xu Qiang.2009.Effects of supplemental different light qualities on growth,photosynthesis,biomass partition and early yield of greenhouse cucumber.Scientia Agricultura Sinica,42(7):2615-2623.(in Chinese)倪纪恒,陈学好,陈春宏,徐强.2009.补充不同光质对温室黄瓜生长发育光合和期产量的影响.中国农业科学,42(7):2615-2623.
Ni W,Xu S L,Tepperman J M,Stanley D J,Maltby D A,Gross J D,Burlingame A L,Wang Z Y,Quail P H.2014.A mutually assured destruction mechanism attenuates light signaling in Arabidopsis.Science,344(6188):1160-1164.
Ni Z,Kim E D,Ha M,Lackey E,Liu J,Zhang Y,Sun Q,Chen Z J.2009.Altered circadian rhythms regulate growth vigour in hybrids and allopolyploids.Nature,457(7277):327-331.
Noren L,Kindgren P,Stachula P,Ruhl M,Eriksson M E,Hurry V,Strand A.2016.Circadian and plastid signaling pathways are integrated to ensure correct expression of the CBF and COR genes during photoperiodic growth.Plant Physiology,171(2):1392-1406.
Nyitrai P,Sarvari E,Keresztes A,Lang F.1994.Organization of thylakoid membranes in low light grown maize seedlings.Effect of lincomycin treatment.Journal of Plant Physiology,144(3):370-375.
Oda-Yamamizo C,Mitsuda N,Sakamoto S,Ogawa D,Ohme-Takagi M,Ohmiya A.2016.The NAC transcription factor ANAC046 is a positive regulator of chlorophyll degradation and senescence in Arabidopsis leaves.Scientific Reports,6:23609.
Osterlund M T,Hardtke C S,Wei N,Deng X W.2000.Targeted destabilization of HY5 during lightregulated development of Arabidopsis.Nature,405(6785):462-466.
Ouyang X,Li J,Li G,Li B,Chen B,Shen H,Huang X,Mo X,Wan X,Lin R,Li S,Wang H,Deng X W.2011.Genome-wide binding site analysis of FAR-RED ELONGATED HYPOCOTYL3 reveals its novel function in Arabidopsis development.The Plant Cell,23(7):2514-2535.
Papenbrock J,Mock H P,Kruse E,Grimm B.1999.Expression studies in tetrapyrrole biosynthesis:inverse maxima of magnesium chelatase and ferrochelatase activity during cyclic photoperiods.Planta,208(2):264-273.
Park J E,Jeong B R,Seungjae H.2013.Growth of lettuce in closed-type plant production system as affected by light intensity and photoperiod under influence of white LED light.Protected Horticulture and Plant Factory,22(3):228-233.
Pattanayak G K,Tripathy B C.2011.Overexpression of protochlorophyllide oxidoreductase C regulates oxidative stress in Arabidopsis.PLoS ONE,6(10):e26532.
Petroutsos D,Tokutsu R,Maruyama S,Flori S,Greiner A,Magneschi L.,Cusant L,Kottke T,Mittag M,Hegemann P,Finazzi G,Minagawa J.2016.A blue-light photoreceptor mediates the feedback regulation of photosynthesis.Nature,537(7621):563-566.
Powell A L,Nguyen C V,Hill T,Cheng K L,Figueroa-Balderas R,Aktas H,Ashrafi H,Pons C,Fernández-Mu?oz R,Vicente A,Lopez-Baltazar J,Barry C S,Liu Y,Chetelat R,Granell A,Van Deynze A,Giovannoni J J,Bennett A B.2012.Uniform ripening encodes a Golden 2-like transcription factor regulating tomato fruit chloroplast development.Science,336(6039):1711-1715.
Qiu K,Li Z,Yang Z P,Chen J Y,Wu S X,Zhu X Y,Gao S,Gao J,Ren G D,Kuai B,Zhou X.2015.EIN3 and ORE1 accelerate degreening during ethylene-mediated leaf senescence by directly activating chlorophyll catabolic genes in Arabidopsis.PLoS Genetics,11(7):e1005399.
Reid J D,Hunter C N.2004.Magnesium-dependent ATPase activity and cooperativity of magnesium chelatase from Synechocystis sp.PCC6803.Journal of Biological Chemistry,279(26):26893-26899.
Rosado D,Gramegna G,Cruz A,Lira B S,Freschi L,de Setta N,Rossi M.2016.Phytochrome interacting factors(PIFs)in Solanum lycopersicum:diversity,evolutionary history and expression profiling during different developmental processes.PLoS ONE,11(11):e0165929.
Rusaczonek A,Czarnocka W,Kacprzak S,WitońD,?lesak I,Szechyńska-Hebda M,Gawroński P,Karpiński S.2015.Role of phytochromes A and B in the regulation of cell death and acclimatory responses to UR stress in Arabidopsis thaliana.Journal of Experimental Botany,66(21):6679-6695.
Saebo A,Krekling T,Appelgren M.1995.Light quality affects photosynthesis and leaf anatomy of birch plantlets in vitro.Plant Cell Tissue&Organ Culture,41(2):177-185.
Sakuraba Y,Han S H,Lee S H,H?rtensteiner S,Paek N C.2016.Arabidopsis NAC016 promotes chlorophyll breakdown by directly upregulating STAYGREEN1 transcription.Plant Cell Reports,35(1):155-166.
Sakuraba Y,Jeong J,Kang M Y,Kim J,Paek N C,Choi G.2014.Phytochrome-interacting transcription factors PIF4 and PIF5 induce leaf senescence in Arabidopsis.Nature Communications,14(5):4636.
Sakuraba Y,Kim E Y,Han S H,Piao W,An G,Todaka D,Yamaguchi-Shinozaki K,Paek N C.2017.Rice Phytochrome-Interacting Factor-Like1(Os PIL1)is involved in the promotion of chlorophyll biosynthesis through feed-forward regulatory loops.Journal of Experimental Botany,68(15):4103-4114.
Schmid H C,Oster U,Kogel J,Lenz S,Rudiger W.2001.Cloning and characterisation of chlorophyll synthase from Avena sativa.Biological Chemistry,382(6):903-911.
Servet C,Conde e Silva N,Zhou D X.2010.Histone acetyl-transferase AtGCN5/HAG1 is a versatile regulator of developmental and inducible gene expression in Arabidopsis.Molecular Plant,3(4):670-677.
Shin J,Kim K,Kang H,Zulfugarov I S,Bae G,Lee C H,Lee D,Choi G.2009.Phytochromes promote seedling light responses by inhibiting four negatively-acting phytochrome-interacting factors.Proceedings of the National Academy of Sciences of the United States of America,106(18):7660-7665.
Shi Qing-hua,Zhu Zhu-jun,Ying Quan-shen,Qian Qiong-qiu.2005.Effects of excess Mn on photosynthesis characteristics in cucumber under different light intensity.Chinese Journal of Applied Ecology,16(6):1047-1050.(in Chinese)史庆华,朱祝军,应泉盛,钱琼秋.2005.不同光强下高锰对黄瓜光合作用特性的影响.应用生态学报,16(6):1047-1050.
Song Y,Yang C,Gao S,Zhang W,Li L,Kuai B.2014.Age-triggered and dark-induced leaf senescence require the bHLH transcription factors PIF3,4,and 5.Molecular Plant,7(12):1776-1787.
Stenbaek A,Jensen P E.2010.Redox regulation of chlorophyll biosynthesis.Phytochemistry,71(8-9):853-859.
Stephenson P G,Terry M J.2008.Light signalling pathways regulating the Mg-chelatase branchpoint of chlorophyll synthesis during de-etiolation in Arabidopsis thaliana.Photochemical&Photobiological Sciences,7(10):1243-1252.
Sui X L,Mao S L,Wang L H,Zhang B X,Zhang Z X.2012.Effect of low light on the characteristics of photosynthesis and chlorophyll a fluorescence during leaf development of sweet pepper.Journal of Integrative Agriculture,11(10):1633-1643.
Sun X,Feng P,Xu X,Guo H,Ma J,Chi W,Lin R,Lu C,Zhang L.2011.A chloroplast envelope-bound PHD transcription factor mediates chloroplast signals to the nucleus.Nature Communications,(2):477.
Tang W,Wang W,Chen D,Ji Q,Jing Y,Wang H,Lin R.2012.Transposase-derived proteins FHY3/FAR1 interact with PHYTOCHROME-INTERACTING FACTOR1 to regulate chlorophyll biosynthesis by modulating HEMB1 during deetiolation in Arabidopsis.The Plant Cell,24(5):1984-2000.
Thum K E,Kim M Y,Christopher D A,Mullet J E.2001.Cryptochrome 1,cryptochrome 2,and phytochrome A co-activate the chloroplast psbDblue light-responsive promoter.The Plant Cell,13(12):2747-2760.
Toledo-Ortiz G,Johansson H,Lee K P,Bou-Torrent J,Stewart K,Steel G,Rodríguez-Concepción M,Halliday K J.2014.The HY5-PIF regulatory module coordinates light and temperature control of photosynthetic gene transcription.PLoS Genetics,10(6):e1004416.
Tucker D J.1981.Phytochrome regulation of leaf senescence in cucumber and tomato.Plant Science Letters,23(1):103-108.
Tzvetkova-Chevolleau T,Franck F,Alawady A E,Dall'Osto L,Carrière F,Bassi R,Grimm B,Nussaume L,Havaux M.2007.The light stress-induced protein ELIP2 is a regulator of chlorophyll synthesis.The Plant Journal,50(5):795-809.
Ueda Minoru,Tanaka Ayumi,Sugimoto Kazuhiko,Shikanai Toshiharu,Nishimura Yoshiki.2014.ChlB requirement for chlorophyll biosynthesis under short photoperiod in Marchantia polymorpha L.Genome Biology Evolution,6(3):620-628.
Valdés A E,Rizzardi K,Johannesson H,Para A,Sund?s-Larsson A,Landberg K.2012.Arabidopsis thaliana TERMINAL FLOWER2 is involved in light-controlled signalling during seedling photomorphogenesis.Plant Cell and Environment,35(6):1013-1025.
Vothknecht U C,Kannangara C G,Von W D.1988.Barley glutamyl tRNA-Glu reductase:mutations affecting haem inhibition and enzyme activity.Phytochemistry,47(4):513-519.
Wang Z H,Hong X,Hu K K,Wang Y,Wang X X,Du S Y,Li Y,Hu D D,Cheng K,An B G,Li Y S.2017.Impaired magnesium protoporphyrin IX methyltransferase(ChlM)impedes chlorophyll synthesis and plant growth in rice.Frontiers in Plant Science,8:1-18.
Waters M T,Wang P,Korkaric M,Capper R G,Saunders N J,Langdale J A.2009.GLK transcription factors coordinate expression of the photosynthetic apparatus in Arabidopsis.The Plant Cell,21(4):1109-1128.
Xu G,Guo H,Zhang D,Chen D,Jiang Z,Lin R.2015.REVEILLE1 promotes NADPH:protochlorophyllide oxidoreductase A expression and seedling greening in Arabidopsis.Photosynthesis Research,126(2-3):331-340.
Xu Kai,Guo Yan-ping,Zang Shang-long.2005.Effect of light quality on photosynthesis and chlorophyll fluorescence in strawberry leaves.Scientia Agricultura Sinica,38(2):369-375.(in Chinese)徐凯,郭延平,张上隆.2005.不同光质对草莓叶片光合作用和叶绿素荧光的影响.中国农业科学,38(2):369-375.
Xu Li,Wei Hui,Qi Lian-dong,Hao Shu-qin,Liang Qing-ling,Yu Wen-yan,Liu Shi-qi.2010.Effects of different light qualities on growth and quality of Lactuca sativa L.China Fruit&Vegetable,(4):19-22.(in Chinese)许莉,尉辉,齐连东,郝树芹,梁庆玲,于文艳,刘世琦.2010.不同光质对叶用莴苣生长和品质的影响.中国果菜,(4):19-22.
Xu X,Chi W,Sun X,Feng P,Guo H,Li J,Lin R,Lu C,Wang H,Leister D,Zhang L.2016.Convergence of light and chloroplast signals for de-etiolation through ABI4-HY5 and COP1.Nature Plants,2(6):16066.
Xu X,Hotta C T,Dodd A N,Love J,Sharrock R,Lee Y W,Xie Q G,Johnson C H,Webb A A.2007.Distinct light and clock modulation of cytosolic free Ca2+oscillations and rhythmic CHLOROPHYLL A/B BINDING PROTEIN2 promoter activity in Arabidopsis.The Plant Cell,19(11):3474-3490.
Yang L,Lou Y,Peng Z,Zhao H,Sun H,Gao Z.2015.Molecular characterization and primary functional analysis of PeMPEC,a magnesium-protoporphyrin IX monomethyl ester cyclase gene of bamboo(Phyllostachys edulis).Plant Cell Reports,34(11):2001-2011.
Yang Xiao-jian,Liu Shi-qi,Zhang Zi-kun,Liu Zhong-liang,Ma Lin,Zhang Yu.2011.Effects of different light emitting diode sources on growth and chlorophyll fluorescence parameters in garlic seedling.China Vegetables,(6):62-67.(in Chinese)杨晓健,刘世琦,张自坤,刘中良,马琳,张宇.2011.不同LED光源对青蒜苗生长及叶绿素荧光特性的影响.中国蔬菜,(6):62-67.
Yin X R,Xie X L,Xia X J,Yu J Q,Ferguson I B,Giovannoni J J,Chen K S.2016.Involvement of an ethylene response factor in chlorophyll degradation during citrus fruit degreening.The Plant Journal,86(5):403-412.
Yuan M,Yuan S,Zhang Z W,Xu F,Chen Y E,Du J B,Lin H H.2010.Putative mutation mechanism and light responses of a protochlorophyllide oxidoreductase-less barley mutant NYB.Plant&Cell Physiology,51(8):1361-1371.
Yuan M,Zhang D W,Zhang Z W,Chen Y E,Yuan S,Guo Y R,Lin H H.2012.Assembly of NADPH:protochlorophyllide oxidoreductase complex is needed for effective greening of barley seedlings.Journal of Plant Physiology,169(13):1311-1316.
Yuan S,Zhang Z W,Zheng C,Zhao Z Y,Wang Y,Feng L Y,Niu G,Wang C Q,Wang J H,Feng H,Xu F,Bao F,Hu Y,Cao Y,Ma L,Wang H,Kong D D,Xiao W,Lin H H,He Y.2016.Arabidopsis cryptochrome 1 functions in nitrogen regulation of flowering.Proceedings of the National Academy of Sciences of the United States of America,113(27):7661-7666.
Yuan Y J,Mei L H,Wu M B,Wei W,Shan W,Gong Z H,Zhang Q,Yang F Q,Yan F,Zhang Q,Luo Y Q,Xu X,Zhang W F,Miao MJ,Lu W J,Li Z G,Deng W.2018.SlARF10,an auxin response factor,is involved in chlorophyll and sugar accumulation during tomato fruit development.Journal of Experimental Botany,69(22):5507-5518.
Zha Lingyan,Liu Wenke.2017.Effects of light quality and photoperiod on growth and yield of cherry radish grown under red plus blue LEDs.China Illuminating Engineering Journal,28(6):83-91.(in Chinese)查凌雁,刘文科.2017.LED红蓝光质比及其光周期对樱桃萝卜品种生长与产量的影响.照明工程学报,28(6):83-91.
Zhang D,Li Y,Zhang X,Zha P,Lin R.2017.The SWI2/SNF2 chromatin-remodeling ATPase BRAHMA regulates chlorophyll biosynthesis in Arabidopsis.Molecular Plant,10(1):155-167.
Zhang D W,Yuan S,Xu F,Zhu F,Yuan M,Ye H X,Guo H Q,Lv X,Yin Y,Lin H H.2016.Light intensity affects chlorophyll synthesis during greening process by metabolite signal from mitochondrial alternative oxidase in Arabidopsis.Plant Cell&Environment,39(1):12-25.
Zhang Y,Liu Z,Chen Y,He J X,Bi Y.2015.PHYTOCHROME-INTERACTING FACTOR 5(PIF5)positively regulates dark-induced senescence and chlorophyll degradation in Arabidopsis.Plant Science,237:57-68.
Zhang Z W,Feng L Y,Cheng J,Tang H,Xu F,Zhu F,Zhao Z Y,Yuan M,Chen Y E,Wang J H,Yuan S,Lin H H.2013.The roles of two transcription factors,ABI4 and CBFA,in ABA and plastid signalling and stress responses.Plant Molecular Biology,83(4-5):445-458.
Zhao J,Zhou J J,Wang Y Y,Gu J W,Xie X Z.2013.Positive regulation of phytochrome B on chlorophyll biosynthesis and chloroplast development in rice.Rice Science,20(4):243-248.
Zhong S,Zhao M,Shi T,Shi H,An F,Zhao Q,Guo H.2009.EIN3/EIL1 cooperate with PIF1 to prevent photooxidation and to promote greening of Arabidopsis seedlings.Proceedings of the National Academy of Sciences of the United States of America,106(50):21431-21436.
Zhu X,Chen J,Xie Z,Gao J,Ren G,Gao S,Zhou X,Kuai B.2015.Jasmonic acid promotes degreening via MYC2/3/4-and ANAC019/055/072-mediated regulation of major chlorophyll catabolic genes.The Plant Journal,84(3):597-610.