王峰，王秀杰，趙勝男，閆家榕，卜鑫，張穎，劉玉鳳，許濤，齊明芳，齊紅巖，李天來

光對園藝植物花青素生物合成的調(diào)控作用

王峰，王秀杰，趙勝男，閆家榕，卜鑫，張穎，劉玉鳳，許濤，齊明芳，齊紅巖，李天來

（沈陽農(nóng)業(yè)大學(xué)園藝學(xué)院/設(shè)施園藝省部共建教育部重點實驗室/北方園藝設(shè)施設(shè)計與應(yīng)用技術(shù)國家地方聯(lián)合工程研究中心（遼寧），沈陽 110866）

花青素是植物中一類重要的類黃酮化合物，在植物花朵、果實等器官色澤形成和抗氧化過程中起著重要作用。植物組織中花青素的形成依賴于光信號，但是光信號對花青素生物合成的調(diào)控機(jī)制及信號網(wǎng)絡(luò)很大程度上還不清晰。本文簡述了花青素生物合成及運(yùn)轉(zhuǎn)過程的研究進(jìn)展，簡要歸納了MYB、bHLH、WDR三類主要因子對花青素合成的轉(zhuǎn)錄調(diào)控作用，重點闡釋光信號（光強(qiáng)、光質(zhì)、光照時長）對植物花青素合成的調(diào)控作用。研究表明，光環(huán)境（光強(qiáng)、光質(zhì)、光照時長）主要通過不同的光受體（UVR8、CRYs、PHOTs、PHYs）影響光信號通路重要因子COP1的泛素化能力和HY5的穩(wěn)定性，以及其他光信號轉(zhuǎn)錄因子如光敏色素互作因子PIFs的穩(wěn)定性，進(jìn)而調(diào)控花青素的生物合成過程。這些光信號因子一方面直接結(jié)合到調(diào)控花青素合成的MYB、bHLH、WDR三大類轉(zhuǎn)錄因子上，轉(zhuǎn)錄激活或抑制它們的表達(dá)進(jìn)而調(diào)控花青素的合成；另一方面，這些光信號因子通過與MYB、bHLH、WDR三大類轉(zhuǎn)錄因子蛋白互作，影響它們形成的MBW復(fù)合體穩(wěn)定性，進(jìn)而調(diào)控花青素的合成。此外，這些光信號因子還可以通過不依賴于MBW復(fù)合體的通路調(diào)控花青素的合成，如HY5通過調(diào)控影響花青素的生物合成；另外，一些未知的光響應(yīng)因子可能以不依賴MBW通路的方式直接或間接地調(diào)控花青素合成基因和液泡膜上的運(yùn)轉(zhuǎn)蛋白，改變液泡酸化，調(diào)節(jié)花青素的合成。同時，光信號會影響光合電子傳遞，光合電子傳遞鏈中的一些因子也會通過依賴和不依賴MBW的途徑影響植物花青素的合成。這些途徑如何協(xié)調(diào)以及哪些信號因子優(yōu)先受光環(huán)境（光強(qiáng)、光質(zhì)、光照時間）調(diào)控？本文為深入研究光信號對花青素生物合成的調(diào)控機(jī)理提供參考，以探索光調(diào)控花青素積累的有效途徑及靶標(biāo)分子，為利用基因工程、代謝工程和光環(huán)境調(diào)控手段改良園藝植物花青素積累提供理論基礎(chǔ)。

光；花青素；轉(zhuǎn)錄因子；轉(zhuǎn)錄調(diào)控；園藝植物

花青素是自然界中廣泛存在于植物中的水溶性天然色素，在植物的花、葉片和果實等器官中均有積累?；ㄇ嗨乜梢酝ㄟ^其艷麗的色彩幫助植物吸引昆蟲和飛鳥等媒介，幫助植物傳粉和散播種子。另外，花青素具有較強(qiáng)的抗氧化能力，可以減輕植物組織免受活性氧（ROS）等脅迫的傷害，同時，在抗衰老[1]及防治心血管疾病[2]等方面對人體也有保健功效。光環(huán)境作為影響植物花青素合成的重要環(huán)境因子，受到越來越多的關(guān)注，解析光對花青素生物合成的調(diào)控作用，對精準(zhǔn)調(diào)控園藝植物不同器官花青素的含量、提高園藝產(chǎn)品營養(yǎng)成分及人體保健等方面具有重要的研究和應(yīng)用價值。

1 花青素的生物合成代謝

花青素生物合成途徑是植物類黃酮途徑的一個分支途徑，主要在內(nèi)質(zhì)網(wǎng)表面進(jìn)行。一般來說，花青素由3個丙二酰輔酶A分子和一個4-香豆酰輔酶A分子在細(xì)胞質(zhì)中結(jié)合，經(jīng)由查爾酮合酶（CHS）作用產(chǎn)生查爾酮（圖1）。查爾酮經(jīng)查爾酮異構(gòu)酶（CHI）催化形成黃烷酮柚皮素，黃烷酮-3-羥化酶（F3H）催化黃烷酮柚皮素形成二氫黃酮醇（DHK）。DHK一方面直接被二氫黃酮醇還原酶（DFR）催化形成無色天竺葵素；另一方面，DHK經(jīng)由二氫黃酮醇-3′-羥化酶（F3′H）作用形成二氫槲皮素（DHQ），DHQ和DHK在二氫黃酮醇-3′, 5′-羥化酶（F3′5′H）作用下形成二氫楊梅素（DHM）。DHQ和DHM被DFR催化后分別形成無色矢車菊素和無色飛燕草素。3種無色花青素在花青素合成酶（ANS）催化下形成天竺葵素、矢車菊素和飛燕草素。不穩(wěn)定的花青素通過類黃酮-3-O-葡萄糖基轉(zhuǎn)移酶（UFGT）催化形成穩(wěn)定的花青苷。矢車菊素苷經(jīng)由甲基轉(zhuǎn)移酶（OMT）催化形成芍藥花苷，飛燕草素苷經(jīng)由OMT催化形成牽?；ㄋ剀蘸湾\葵素苷（圖1）。

花青素苷合成后需要在轉(zhuǎn)運(yùn)蛋白和轉(zhuǎn)運(yùn)囊泡的協(xié)助下有效地向液泡中轉(zhuǎn)運(yùn)并貯藏（圖1），從而防止花青素苷自身被氧化變性及對細(xì)胞造成毒害。由于花青素苷最終所處的液泡環(huán)境會影響花青素苷的呈色，如酸性環(huán)境會使其顏色紅移，而堿性環(huán)境會使其顏色藍(lán)移[3]，因此，明確花青素苷的液泡運(yùn)轉(zhuǎn)機(jī)制極其重要。研究發(fā)現(xiàn)，花青素苷液泡轉(zhuǎn)運(yùn)的主要方式為：在GST協(xié)助下被靶向定位到液泡附近，液泡膜上的MRP類轉(zhuǎn)運(yùn)蛋白識別后將其跨膜轉(zhuǎn)運(yùn)至液泡[4]；由液泡膜上的MATE類轉(zhuǎn)運(yùn)蛋白將其跨膜轉(zhuǎn)運(yùn)到液泡中，這個過程需要如H+-ATPase和H+-pyrophosphatase（V-PPase）質(zhì)子泵將質(zhì)子運(yùn)輸?shù)揭号葜袑σ号菟峄痆5]；由囊泡包裹，細(xì)胞膜導(dǎo)出的囊泡以膜融合的方式直接將花青素運(yùn)輸?shù)劫A藏液泡中，這個過程不依賴GST和MRP3蛋白的協(xié)助[4]。另外，ABC運(yùn)轉(zhuǎn)蛋白也參與花青素苷的液泡吸收和外排過程[6]。

CHS：查爾酮合酶Chalcone synthase；CHI：查爾酮異構(gòu)酶Chalcone isomerase；F3H：黃烷酮-3β-羥化酶Flavanone-3β-hydroxylase；F3′H：二氫黃酮醇-3′-羥化酶Dihydroflavonoid-3'-hydroxylase；F3′5′H：二氫黃酮醇-3′, 5′-羥化酶Dihydroflavonoid-3', 5'-hydroxylase；DFR：二氫黃酮醇還原酶Dihydroflavonol reductase；ANS：花青素合成酶Anthocyanin synthase；UFGT：尿苷二磷酸-葡萄糖-類黃酮-3-葡糖基轉(zhuǎn)移酶UDP-glucose flavonoid 3-glucosyltransferase；OMT：甲基轉(zhuǎn)移酶O-methyltransferases；GST：谷胱甘肽轉(zhuǎn)移酶Glutathione S-transferase；MRP：多藥耐藥相關(guān)蛋白Multidrug resistance-associated protein；MATE：多藥和有毒化合物排出家族蛋白Multidrug and toxic compound extrusion；ABC：C型的ATP結(jié)合蛋白C type of ATP-binding cassette；AVIs：花青素苷液泡內(nèi)涵體Anthocyanic vacuolar inclusions；Phenylalanine：苯基丙氨酸；3×Malonyl CoA：3×丙二酰輔酶A；4-Coumaroyl CoA：4-香豆酰輔酶A；Chalcone：查爾酮；Flavanones：黃烷酮；Dihydrokaempferol（DHK）：二氫黃烷酮；Dihydroquercetin（DHQ）：二氫櫟精；Dihydromyricetin（DHM）：二氫楊梅酮；Leucopelargonidin：無色天竺葵素；Leucocyanidin：無色矢車菊素；Leucodelphinidin：無色飛燕草素；Pelargonidin：天竺葵素；Cyanidin：矢車菊素；Delphinidin：飛燕草素；Pelargonidin 3-glucoside：天竺葵素苷；Cyanidin 3-glucoside：矢車菊素苷；Delphinidin 3-glucoside：飛燕草素苷；Peonidin：芍藥花青素苷；Petunidin：牽?；ㄋ剀?；Malvidin：錦葵素苷；Anthocyanins：花青素；Cytosolic：細(xì)胞質(zhì)；Endoplasmic reticulum：內(nèi)質(zhì)網(wǎng)

2 花青素生物合成的轉(zhuǎn)錄調(diào)控

花青素生物合成途徑的主要基因在轉(zhuǎn)錄水平上受多種轉(zhuǎn)錄因子的調(diào)控，主要包括MYB、bHLH和WDR轉(zhuǎn)錄因子，這3類轉(zhuǎn)錄因子可以形成MBW復(fù)合體[7]。擬南芥中花青素合成基因分為早期合成基因（early biosynthetic genes，EBGs）和后期合成基因（later biosynthetic genes，LBGs），EBGs包括、和′，LBGs包括和[8]。EBGs的表達(dá)不受MBW復(fù)合物調(diào)控，而LBGs的表達(dá)受MBW復(fù)合物調(diào)控[9]。

2.1 MYB轉(zhuǎn)錄因子

MYB蛋白家族N端含有可與DNA結(jié)合的保守MYB結(jié)構(gòu)域，C端在不同物種間序列變異較大，但常有轉(zhuǎn)錄激活或抑制結(jié)構(gòu)域的作用[10]。MYB結(jié)構(gòu)域由1–4個R基序（R repeat）組成，根據(jù)含有R基序的數(shù)量分為1R-MYB、R2R3-MYB、3R-MYB和4R-MYB。其中有2個R基序的R2R3-MYB是最常見的MYB轉(zhuǎn)錄因子，也是參與調(diào)控類黃酮代謝和花青素生物合成的重要轉(zhuǎn)錄因子。MYB蛋白功能異常或靶基因啟動子上MYB識別序列[MYB-recognizing elements，MRE；識別位點ANCNN(C/A)C]異常，都可能導(dǎo)致花青素的生物合成不正常[11]。

玉米中C1（Colorless-1）是植物中首個被發(fā)現(xiàn)的R2R3-MYB轉(zhuǎn)錄因子，它正調(diào)控玉米中和的表達(dá)，從而促進(jìn)玉米花青素的合成[12]。矮牽牛中MYB型轉(zhuǎn)錄因子PhAN2（anthocyanin 2）的C端具有與玉米C1相似的結(jié)構(gòu)域，與PhAN4均正調(diào)控的表達(dá)，從而促進(jìn)矮牽?；ㄇ嗨氐暮铣蒣13]。與矮牽牛PhAN2同源的亞洲雜交百合的LhMYB6和LhMYB12同樣可以促進(jìn)白百合花青素的積累[14]。最近研究發(fā)現(xiàn)，野生番茄SlAN2-like轉(zhuǎn)錄因子可以激活的表達(dá)，誘導(dǎo)番茄果皮中花青素的積累[15]，但SlAN2-like轉(zhuǎn)錄激活的會反饋抑制的表達(dá)[16]；另外，SlAN2-like的選擇性剪切對番茄花青素的積累也有影響[17]。研究發(fā)現(xiàn)，擬南芥MYB轉(zhuǎn)錄因子AtPAP1（production of anthocyanin pigment 1）不僅正調(diào)控和等基因的表達(dá)，而且促進(jìn)糖基轉(zhuǎn)移酶基因和的表達(dá)，誘導(dǎo)矢車菊素大量積累[18]。豆科植物紫花苜蓿、蒺藜苜蓿和白三葉中與AtPAP1同源的LAP1（legume anthocyanin production 1）同樣能夠有效地促進(jìn)矢車菊素積累[19]。蘋果MdMYB10與AtPAP1有較高同源性[20]，MdMYB10結(jié)合到自身啟動子上游23 bp的串聯(lián)重復(fù)序列上，激活自身轉(zhuǎn)錄，進(jìn)而促進(jìn)的表達(dá)，使蘋果果肉花青素積累，而白色果肉蘋果中啟動子上則缺少這段序列，導(dǎo)致的轉(zhuǎn)錄水平較低，破壞了花青素的積累[21]。研究發(fā)現(xiàn)，MYB10在果實花青素形成中是保守的，草莓[22]、甜櫻桃[23]、油桃[24]和梨[25]等果實花青素形成均與MYB10相關(guān)。此外，植物中一些各異的MYB也能夠影響花青素的積累。如擬南芥AtMYB113和AtMYB114[9]，龍膽中的GtMYB3[26]，菊花中的CmMYB6[27]，葡萄中的VvMYBA、VvMYB5a和VvMYB5b[28]，蘋果的MdMYB9、MdMYB11和MdMYB110[29-30]，以及番茄SlMYB75[31]。

為保證花青素在植物體內(nèi)的代謝平衡，一些MYB在花青素生物合成中起著負(fù)調(diào)控因子的作用。如擬南芥AtMYB2負(fù)調(diào)控、和的表達(dá)[32]；葡萄VvMYB4-like抑制和的表達(dá)[33]；蘋果MdMYB6過表達(dá)會抑制和等基因的表達(dá)[34]；智利草莓FcMYB1會抑制的表達(dá)[35]。這些抑制型MYB一方面直接轉(zhuǎn)錄抑制花青素合成途徑相關(guān)基因的表達(dá)；另一方面通過與調(diào)控花青素合成的正調(diào)控因子結(jié)合，抑制它們的表達(dá)或破壞MBW復(fù)合體的形成。如具有獨特TLLLFR抑制結(jié)構(gòu)域的AtMYBL2轉(zhuǎn)錄因子，通過與AtTT8結(jié)合進(jìn)而抑制和的表達(dá)[36]；AtSPL9（squamosa promoter binding protein-like 9）轉(zhuǎn)錄因子通過破壞MBW復(fù)合體的穩(wěn)定性抑制的表達(dá)[37]；葡萄中的VvMYBC2-L1和VvMYBC2-L3與VvAN1結(jié)合，破壞MBW復(fù)合體的穩(wěn)定性，抑制花青素的積累[38]。研究發(fā)現(xiàn)，矮牽牛PhMYB27一方面通過與PhAN1互作，激活其C末端EAR基序抑制的表達(dá)[39]；另一方面通過與bHLH型轉(zhuǎn)錄因子PhJAF13和PhAN11結(jié)合互作，破壞MBW復(fù)合體形成，使MBW復(fù)合物從激活狀態(tài)變?yōu)橐种茽顟B(tài)，進(jìn)而抑制花青素合成[40]。AtCPC（CAPRICE）與PhMYB27不同，它是C末端不含抑制基序的R3-MYB，不能直接與MBW復(fù)合物結(jié)合，而是間接減少MBW復(fù)合物的活性，從而抑制花青素的生物合成[41]。

2.2 bHLH轉(zhuǎn)錄因子

bHLH（basic helix-loop-helix）轉(zhuǎn)錄因子具有與DNA結(jié)合的高度保守的bHLH結(jié)構(gòu)域，這個結(jié)構(gòu)域的N端由13—18個親水氨基酸殘基組成堿性區(qū)（basic），負(fù)責(zé)與DNA接觸，C端為HLH域，主要參與同源或異源二聚體的形成，其中的環(huán)狀區(qū)（Loop，L）連接著2個螺旋（helix，H）。bHLH轉(zhuǎn)錄因子決定了MBW復(fù)合物識別靶基因啟動子上的轉(zhuǎn)錄結(jié)合位點及激活靶基因轉(zhuǎn)錄的特異性[42]。研究表明，花青素相關(guān)的bHLH對靶基因的有效識別序列（bHLH-recognizing elements，BRE）為CACN(A/C/T)(G/T)[11]。bHLH蛋白功能異?；虬谢騿幼由螧RE異常，都可能導(dǎo)致花青素合成異常。

最早發(fā)現(xiàn)的一批調(diào)控花青素合成的bHLH家族轉(zhuǎn)錄因子是玉米中的R1、B1、Lc（Leaf color）和Sn[43-44]。ZmLc能夠激活和的表達(dá)，促進(jìn)花青素的生物合成[13]。與玉米同源的擬南芥bHLH轉(zhuǎn)錄因子為AtTT8（transparent testa 8）、AtGL3（glabra 3）和AtEGL3（enhancer of glabra 3），它們通過調(diào)控的表達(dá)來促進(jìn)花青素的生物合成[45]。蒺藜苜蓿MtTT8是與擬南芥AtTT8同源的bHLH型轉(zhuǎn)錄因子，它能與MtLAP1和MtWD40-1互作形成MBW復(fù)合體，促進(jìn)花青素的合成[46]。研究發(fā)現(xiàn)，矮牽牛中PhJAF13和PhAN1是調(diào)控花青素生物合成的主要bHLH類調(diào)控因子，它們均能與PhAN2互作激活的表達(dá)，且PhAN1還可以直接激活的表達(dá)[42]。同樣，煙草中的NtAN1a與NtAN2互作激活和的表達(dá)[47]。非洲菊bHLH調(diào)控因子GMYC1與MYB轉(zhuǎn)錄因子AN2和GMYB10互作，特異地在花冠和心皮部位促進(jìn)的表達(dá)[48]；紫色天葵的bHLH型基因和MYB轉(zhuǎn)錄因子GbMYB1共同表達(dá)能激活和啟動子，誘導(dǎo)花青素合成[49]。另外，蘋果MdbHLH3和MdbHLH33共同與MdMYB10轉(zhuǎn)錄因子互作，促進(jìn)果實顏色變紅[20]；荔枝LcbHLH1 和LcbHLH3 轉(zhuǎn)錄因子與LcMYB1共同作用，促進(jìn)花青素積累[50]；龍膽GtbHLH1蛋白能與GtMYB3互作，促進(jìn)龍膽花瓣花青素合成[26]；亞洲百合LhbHLH2能與LhMYB6和LhMYB12相互作用，激活、、的表達(dá)促進(jìn)花青素的合成[14]。以上結(jié)果表明，bHLH轉(zhuǎn)錄因子除了直接調(diào)控的表達(dá)外，還可以與MYB轉(zhuǎn)錄因子互作，調(diào)控花青素的合成。

2.3 WDR蛋白家族

WDR（WD40 repeat proteins，WDR）蛋白家族具有保守而特異的二肽重復(fù)基序，每個重復(fù)的WD基元大概有40—60個氨基酸殘基組成，WDR在蛋白質(zhì)互作時作為支架起固定作用。矮牽牛的PhAN11是第1個被發(fā)現(xiàn)調(diào)控花青素生物合成的WDR蛋白，它作用于PhAN2的上游，通過激活的表達(dá)調(diào)控花青素的合成[51]。擬南芥AtTTG1（transparent testa glabra 1）與矮牽牛PhAN11同源，它一方面與bHLH型轉(zhuǎn)錄因子AtGL3蛋白互作，直接促進(jìn)及的表達(dá)；另一方面，AtTTG1與AtPAP1及AtGL3/AtEGL3互作，形成MYB/bHLH/TTG1轉(zhuǎn)錄復(fù)合體，進(jìn)而調(diào)控花青素的生物合成[9]。玉米中約有20個基因影響其花青素的產(chǎn)生，其中ZmPAC1（pale aleurone color 1）是類似于矮牽牛PhAN11和擬南芥AtTTGl的WDR蛋白，的缺失導(dǎo)致花青素含量降低[52]。研究發(fā)現(xiàn)，楊梅MrWD40-1與MrMYB1和MrbHLH1相互作用，參與楊梅花青素的積累[53]；蘋果MdTTG1與bHLH轉(zhuǎn)錄因子和MYB 轉(zhuǎn)錄因子相互作用形成復(fù)合物，激活花青素合成基因和的轉(zhuǎn)錄[54]；藍(lán)莓WD40轉(zhuǎn)錄因子VcWDL2與VcMYBL1和VcbHLHL1互作，參與調(diào)控藍(lán)莓果實花青素的合成[55]；此外，辣椒果實中的沉默后，辣椒的花青素含量明顯減少[56]。盡管WDR屬于廣譜性表達(dá)蛋白、組織特異性較弱，但其一般無冗余拷貝，所以它的突變會影響MBW復(fù)合體的形成或復(fù)合體的定位及信號傳遞，最終破壞了MBW復(fù)合體對眾多靶基因的表達(dá)調(diào)控。

2.4 其他類型調(diào)控因子

除MYB、bHLH和WDR外，miRNA、JAZ蛋白、bZIP蛋白等也參與了植物花青素生物合成的調(diào)控。研究發(fā)現(xiàn)，miR828/TAS4-siR81(-)能與花青素正調(diào)控因子PAP1、PAP2、MYB113結(jié)合，抑制它們的表達(dá)，進(jìn)而負(fù)調(diào)控擬南芥花青素的積累[57]。miR156靶向定位的SPL9轉(zhuǎn)錄因子可使MBW轉(zhuǎn)錄復(fù)合體變得不穩(wěn)定，最終負(fù)調(diào)控擬南芥花青素的生物合成[37]。另外，番茄miR858可以抑制R2R3-MYB的表達(dá)，減少花青素的合成[58]。擬南芥茉莉酸信號通路中的AtJAZ蛋白不僅可以與bHLHs（AtGL3、AtEGL3、AtTT8）和R2R3-MYBs（AtPAP1、AtPAP2）蛋白結(jié)合，阻礙MBW復(fù)合體的形成，抑制花青素的生物合成，而且可以作為正調(diào)控因子參與JA誘導(dǎo)的花青素積累[59]。另外，bZIP型轉(zhuǎn)錄因子，如HY5不僅可以直接調(diào)控和等花青素結(jié)構(gòu)基因的表達(dá)[60]，而且可以結(jié)合到、、等轉(zhuǎn)錄因子啟動子上影響它們的表達(dá)，同時還可以通過調(diào)控的表達(dá)，間接調(diào)控花青素相關(guān)基因的表達(dá)[61-63]。此外，HY5還能與PIF3（phytochrome interacting factor 3）等bHLH蛋白互作，共同調(diào)控花青素合成途徑的下游靶基因，從而促進(jìn)花青素的合成[64]。

3 光調(diào)控植物花青素的生物合成

光是影響植物花青素合成的最重要環(huán)境因子之一。在植物綠色組織或細(xì)胞中，光通過光受體及光合電子傳遞調(diào)節(jié)植物花青素的合成與積累，從而保護(hù)植物組織免受活性氧（ROS）等的脅迫及調(diào)控植物色澤的形成。因此，本文對光信號調(diào)控植物花青素合成及代謝進(jìn)行總結(jié)（圖2），具體內(nèi)容如下。

3.1 光強(qiáng)調(diào)控植物花青素的生物合成

強(qiáng)光可以刺激許多植物花青素的形成與積累[65]。番茄（）果實暴露在光下部分比遮陰部分的花青素含量高[66]；非洲菊花序進(jìn)行黑暗處理后，其花青素被抑制[67]。主要原因是光可以提高苯丙氨酸解氨酶（PAL）、查耳酮合酶（CHS）、二氫黃酮醇4-還原酶（DFR）、類黃酮葡萄糖苷轉(zhuǎn)移酶（UFGT）等花青素合成途徑中關(guān)鍵合成酶的活性，進(jìn)而促進(jìn)花青素的合成和積累。研究發(fā)現(xiàn)，強(qiáng)光可以促進(jìn)花青素生物合成途徑結(jié)構(gòu)基因和調(diào)節(jié)基因的表達(dá)。如強(qiáng)光促進(jìn)擬南芥花青素合成結(jié)構(gòu)基因、及調(diào)節(jié)基因的表達(dá)，從而促進(jìn)植物花青素的合成與積累[68]。強(qiáng)光促進(jìn)矮牽牛和的表達(dá)，而弱光或黑暗致使矮牽牛和紫蘇等植物的花青素結(jié)構(gòu)基因表達(dá)量下調(diào)甚至不表達(dá)，使植株出現(xiàn)白花或淺色花[69-70]。草莓果實中和等結(jié)構(gòu)基因及轉(zhuǎn)錄因子FaMYB10、FaMYB1的表達(dá)量會隨著光照強(qiáng)度的降低而降低，弱光會使草莓紅色減弱、花青素含量降低[71]。強(qiáng)光可以誘導(dǎo)番茄和辣椒等植物R2R3-MYB轉(zhuǎn)錄因子SlAN2和CaMYBa的表達(dá)，抑制含抑制基序的矮牽牛的表達(dá)，從而促進(jìn)花青素的積累[39]。另外，強(qiáng)光可以促進(jìn)番茄SlAN11、SlTT8和SlAN2蛋白結(jié)合，形成MBW復(fù)合體，進(jìn)而促進(jìn)番茄花青素結(jié)構(gòu)基因的表達(dá)及花青素的積累，而SlMYBL2則反饋抑制花青素合成基因的表達(dá)[72]。強(qiáng)光可以增加蘋果的表達(dá)，從而誘導(dǎo)其下游基因的表達(dá)，促進(jìn)果皮中花青素的積累[73]。另外，強(qiáng)光可以通過藍(lán)光受體CRY1（crytochrome 1）誘導(dǎo)的表達(dá)，進(jìn)而促進(jìn)植物花青素的積累[74]。此外，光合電子傳遞可能在植物花青素合成中也起著重要的作用[75]，因此，光信號對植物花青素合成的調(diào)控是多通路協(xié)作的復(fù)雜過程。

3.2 光質(zhì)調(diào)控植物花青素的生物合成

3.2.1 紫外光（UV）和藍(lán)光 在植物花青素合成和積累的過程中，不同光質(zhì)對植物花青素形成的調(diào)控作用不同。對于大多數(shù)植物，紫外線（UV）是花朵成色、花青素積累的重要因子[76]。UV按波長由短到長依次分為UV-C、UV-B和UV-A。研究發(fā)現(xiàn)，UV-C可以促進(jìn)紫甘藍(lán)花青素?；D(zhuǎn)移酶基因以及R2R3-MYB家族轉(zhuǎn)錄因子MYB114和PAP1的表達(dá)，進(jìn)而促進(jìn)花青素合成[77]。另外，UV-C可以提高楊梅果實中的PAL、CHI、4-香豆酰輔酶A連接酶（4CL）和肉桂酸氫化酶（C4H）的活性，從而增加花青素和類黃酮物質(zhì)的合成與積累[78]。研究發(fā)現(xiàn)，一定程度的UV-B可以通過加強(qiáng)藍(lán)莓中CHI酶活性提高其花青素的含量[79]。UV-B在植物體內(nèi)主要通過兩種形式誘導(dǎo)花青素的積累。植物UVR8（UV resistance locus 8）感知UV-B輻射后，由二聚體形式轉(zhuǎn)化為單體形式[80]。單體形式UVR8和COP1、SPA聚合產(chǎn)生UVR8-SPA- COP1復(fù)合物[81]，并聚集在細(xì)胞核中。這種復(fù)合體一方面引起MYB、bHLH和WDR 3種轉(zhuǎn)錄因子響應(yīng)，促進(jìn)MBW復(fù)合體形成，直接或間接促進(jìn)花青素合成途徑中各基因的表達(dá)[82]。如UV-B可以誘導(dǎo)萵苣葉片中的表達(dá)[83]，以及甘藍(lán)和的表達(dá)[84]，這可能與UVR8和MYB13的互作相關(guān)[85]。另一方面，UVR8-SPA-COP1復(fù)合物可穩(wěn)定HY5蛋白[86]（圖2），HY5轉(zhuǎn)錄因子可以激活R2R3-MYB轉(zhuǎn)錄因子，促進(jìn)植物花青素合成基因的表達(dá)，從而誘導(dǎo)花青素的積累[61]。如UV-B下，蘋果中MdHY5與的啟動子結(jié)合，激活其轉(zhuǎn)錄，進(jìn)而促進(jìn)蘋果花青素的合成[64]。研究發(fā)現(xiàn)，溫室中生長的茄子由于UV照射較少，果實著色不良，當(dāng)對茄子進(jìn)行UV-A補(bǔ)光后，茄子顏色加深[87]。研究報道，與白光相比，UV-A可以提高番茄幼苗和果實中花青素的積累[88]，同時，UV-A可以誘導(dǎo)蕪菁和的表達(dá)[89]。

植物通過隱花色素CRY感受UV-A和藍(lán)光[90]。研究發(fā)現(xiàn)，擬南芥突變體中表達(dá)下調(diào)，花青素積累減弱[76]。藍(lán)光下，CRY1和CRY2可以與COP1和WD40相互作用，但二者作用機(jī)制不同。CRY1與SPA1（suppressor of phyA1）互作，將SPA1與COP1隔離，從而阻礙COP1-SPA1蛋白復(fù)合體的形成[91]；而CRY2-SPA1互作則增強(qiáng)了CRY2-COP1的互作，CRY1/2-SPA1相互作用均減弱了COP1的E3泛素連接酶活性，從而使COP1下游轉(zhuǎn)錄因子的蛋白處于穩(wěn)定狀態(tài)[92]。研究發(fā)現(xiàn)，CRY1和CRY2可以激活光信號轉(zhuǎn)錄因子HY5和COL5，進(jìn)而促進(jìn)花青素生物合成結(jié)構(gòu)基因及的表達(dá)，促進(jìn)花青素的積累[86]。研究發(fā)現(xiàn)，茄子SmCRYs在光下抑制SmCOP1的活性，促使SmHY5和SmMYB1結(jié)合到和的啟動子上激活它們的表達(dá)，進(jìn)而促進(jìn)茄子花青素的積累；而黑暗下，SmCRYs不能抑制SmCOP1的活性，SmHY5和SmMYB1被SmCOP1靶定并通過26S泛素蛋白酶體途徑降解，阻止了依賴SmMYB1激活的花青素合成途徑[93]。最近研究發(fā)現(xiàn)，擬南芥轉(zhuǎn)錄共激活因子AtAN3（angustifolia 3）可以結(jié)合到的啟動子上，負(fù)調(diào)控的表達(dá)；缺失后，植物的花青素含量明顯降低[94]（圖2），但是否通過光受體影響植物花青素的合成尚不清楚。研究發(fā)現(xiàn)，增加藍(lán)光的比例可以提高番茄幼苗的花青素含量[95]。藍(lán)光促進(jìn)楊梅果實MrMYB1及花青素合成相關(guān)結(jié)構(gòu)基因和的表達(dá)，從而促進(jìn)果實中花青素的合成[96]。櫻桃在藍(lán)光和白-藍(lán)-綠光的照射下增加了PAL酶的活性，使果實中的花青素含量升高[97]。另外，藍(lán)光可以通過PHOT和CRY受體增加草莓中花青素的積累[98-99]。以上結(jié)果表明，短波長的UV和藍(lán)光可以通過UVR8和CRY，在轉(zhuǎn)錄和轉(zhuǎn)錄后方面影響MBW復(fù)合體或調(diào)控下游光信號轉(zhuǎn)錄因子的表達(dá)，進(jìn)而影響植物花青素的生物合成，但其作用機(jī)制還待進(jìn)一步深入研究。

3.2.2 紅光和遠(yuǎn)紅光 紅光和遠(yuǎn)紅光在植物花青素的生物合成中也起著重要作用。研究發(fā)現(xiàn)，紅光誘導(dǎo)茄科蔬菜中的表達(dá)及覆盆子果實中黃酮類物質(zhì)的合成，進(jìn)而促進(jìn)花青素的合成[100]。另外，高紅光/遠(yuǎn)紅光比例下生長的番茄，其花青素正常合成，但低紅光/遠(yuǎn)紅光照射下，番茄花青素的合成卻受到嚴(yán)重抑制[101-102]，這主要是高比例的紅光可以促進(jìn)DFR酶和花色素雙氧酶（LDOX）活性，進(jìn)而促進(jìn)花青素的積累，而遠(yuǎn)紅光增多（低比例的紅光/遠(yuǎn)紅光）抑制DFR酶和LDOX酶的催化作用[101]。光敏色素是感受紅光和遠(yuǎn)紅光的主要光受體。研究發(fā)現(xiàn)，紅光受體phyB主要通過兩種方式發(fā)揮作用（圖2）：紅光使phyB以活性的Pfr形式從細(xì)胞質(zhì)進(jìn)入細(xì)胞核，一方面通過光敏色素-轉(zhuǎn)錄因子途徑直接調(diào)控基因表達(dá)；另一方面通過光敏色素-COP1途徑間接調(diào)控基因表達(dá)[103]。但是遠(yuǎn)紅光信號調(diào)控花青素合成的機(jī)制尚不清楚，因為遠(yuǎn)紅光受體光敏色素A（phyA）和紅光受體phyB均能夠抑制COP1的活性，而COP1負(fù)調(diào)控花青素的合成[104]，但增加遠(yuǎn)紅光的比例（低紅光/遠(yuǎn)紅光比例）卻降低花青素的積累，這可能是由于遠(yuǎn)紅光對phyB的鈍化作用大于遠(yuǎn)紅光對phyA的激活作用。

UVR8：UV-B光受體；CRYs：隱花色素；PHYs：光敏色素；MBW：MYB-bHLH-WDR構(gòu)成的復(fù)合體；COP1：暗形態(tài)建成中的E3泛素連接酶蛋白；HY5：光信號轉(zhuǎn)錄因子；PIFs：光敏色素互作因子；AN3：花青素合成調(diào)控因子；PET：光合電子傳遞；MRE：MYB識別序列；BRE：bHLH識別序列UVR8: UV resistance locus 8; CRYs: crytochrome; PHYs: phytochromes; MBW: MYB-bHLH-WDR complex; COP1: constitutively photomorphogenic 1; HY5: elongated hypocotyl 5; PIFs: phytochrome interacting factors; AN3: ANGUSTIFOLIA3; PET: photosynthetic electron transport; MRE: MYB-recognizing elements; BRE: bHLH-recognizing elements

3.3 光周期（光照時長）對植物花青素生物合成的調(diào)控

光周期也會影響植物花青素的生物合成。研究表明，利用補(bǔ)光技術(shù)延長光照時間能顯著提高植物花青素的積累[105]。光照可以提高煙草花青素生物合成基因的表達(dá)量，增加花青素的合成，而黑暗則作用相反[8]。對蕪菁地下根進(jìn)行光照處理時，表達(dá)量隨著光照時間的延長逐漸增加[106]。另外，延長光照時間顯著增強(qiáng)紫葉李葉片中PAL酶的活性，促進(jìn)花青素的生物合成，使其葉色變紅[107]。黑暗下植物花青素含量較低可能與COP1有關(guān)，因為突變體在黑暗中能產(chǎn)生花青素[104]。黑暗下，COP1在細(xì)胞核中大量積累，它可能通過E3泛素途徑將其下游光信號及花青素正調(diào)控轉(zhuǎn)錄因子降解[92]。另外，黑暗中，SPA下調(diào)PAP1和PAP2轉(zhuǎn)錄水平[108]。因此，COP1-SPA復(fù)合體是光照或黑暗調(diào)控植物花青素生物合成的中心調(diào)節(jié)因子[104]。然而，并非光照時間越長，花青素積累越多。如亞洲雜交百合‘Vivaldi’在黑暗條件下生長時，花青素含量很低，光照后迅速增加，但光照時間過長后，花青素含量又逐漸降低[14]。因此，明確光照時長對植物花青素積累的影響，對合理利用LED光譜技術(shù)調(diào)控植物花青素的積累有重要意義。

光環(huán)境及光受體（UVR8、CRYs、PHOTs、PHYs）主要通過COP1、HY5、PIFs等光信號因子來調(diào)控植物花青素的合成，這些光信號因子一方面直接結(jié)合到調(diào)控花青素合成的MYB、bHLH、WDR三大類轉(zhuǎn)錄因子上（圖2），轉(zhuǎn)錄激活或抑制它們的表達(dá)[61-63]；另一方面，這些光信號因子通過與MYB、bHLH、WDR三大類轉(zhuǎn)錄因子蛋白互作，影響它們形成的MBW復(fù)合體的穩(wěn)定性[93,104]，進(jìn)而調(diào)控花青素的合成。此外，這些光信號因子還可以通過不依賴MBW的通路調(diào)控花青素的合成[64-65]，如HY5通過調(diào)控影響花青素的合成[58,60]；另外，一些未知的光響應(yīng)因子可能以不依賴MBW通路的方式直接或間接地調(diào)控花青素合成基因和液泡膜上的運(yùn)轉(zhuǎn)蛋白，改變液泡酸化，調(diào)節(jié)花青素的合成[5-6]。

4 光環(huán)境和生物工程改良園藝植物的花青素

4.1 通過調(diào)控光環(huán)境改良園藝植物花青素

花青素既影響植物的色澤，也作為抗氧化物質(zhì)對人體健康有利，因此，提高園藝作物花青素含量成為改善果實營養(yǎng)的熱點。光強(qiáng)、光質(zhì)、光照時長等光環(huán)境，可以改變園藝植物花青素的含量。如增加光照可以促進(jìn)蘋果果皮積累花青素，使果皮變紅[109]。番茄（）果實暴露在光下部分比遮陰部分的花青素含量高[66]。強(qiáng)光誘導(dǎo)番茄和辣椒的和表達(dá)，促進(jìn)番茄和辣椒花青素的積累[39]。當(dāng)對茄子進(jìn)行UV-A補(bǔ)光后，茄子顏色加深[87]。增加光環(huán)境中藍(lán)光的比例可以促進(jìn)番茄中花青素的含量[95]。研究表明，光環(huán)境的調(diào)控是改良園藝作物花青素的重要手段，因此，LED補(bǔ)光技術(shù)成為調(diào)控園藝作物色澤的關(guān)鍵技術(shù)。

4.2 通過調(diào)控關(guān)鍵基因進(jìn)行生物工程改良

傳統(tǒng)雜交和誘變?yōu)橹鞯挠N技術(shù)對園藝植物花青素的改良周期長。通過改變調(diào)控花青素關(guān)鍵基因的表達(dá)水平進(jìn)而改變花青素積累，是花青素生物工程和代謝工程的一個重要策略。如金魚草和轉(zhuǎn)入番茄后，轉(zhuǎn)基因番茄的果皮和果肉呈現(xiàn)深紫色，花青素含量遠(yuǎn)高于藍(lán)莓[110]，這樣的番茄對提高人體抗氧化能力等有重要的功效。玫瑰、康乃馨和菊花中過量表達(dá)，花朵合成飛燕草素呈現(xiàn)藍(lán)紫色，解決了傳統(tǒng)育種技術(shù)培育不出富含飛燕草素藍(lán)色花朵的難題[111-112]。將轉(zhuǎn)入煙草中可產(chǎn)生1種具有藥用價值的花青素Cyanidin 3-O-rutinoside，且這種成分占總花青素含量的98%[113]。這一研究使工業(yè)上提取大量具有藥用功能的花青素成為可能，同時以煙草作為原料降低了生產(chǎn)成本，極大地促進(jìn)了花青素代謝工程和生物工程的發(fā)展。

5 展望

近年來，人類比以往更加關(guān)注花青素在植物色澤形成、抗氧化、人體保健等方面的作用，而光作為花青素合成的重要因子也受到越來越多的關(guān)注。解析光對花青素生物合成的調(diào)控網(wǎng)絡(luò)對精準(zhǔn)調(diào)控園藝植物不同器官花青素的含量有重要的生物學(xué)意義。盡管人類利用模式植物（矮牽牛和擬南芥）對花青素的生物合成途徑有了較深的理解，并且陸續(xù)克隆和鑒定了一些其他物種花青素合成和運(yùn)轉(zhuǎn)途徑的基因，但關(guān)于光信號對植物花青素生物合成及運(yùn)轉(zhuǎn)調(diào)控機(jī)制的研究才剛剛起步，仍有許多問題值得進(jìn)一步思考：（1）不同光受體與MBW復(fù)合體的互作調(diào)控關(guān)系尚不清晰。目前除了HY5對花青素積累的調(diào)控機(jī)制取得一定進(jìn)展外，與MBW復(fù)合體互作的其他光信號因子及其功能的解析，對構(gòu)建光信號調(diào)控花青素的信號網(wǎng)絡(luò)意義重大；（2）盡管已經(jīng)發(fā)現(xiàn)了許多調(diào)控花青素生物合成的正調(diào)控因子，但其負(fù)調(diào)控因子研究相對較少，尤其是光對這些負(fù)調(diào)控因子的作用機(jī)制嚴(yán)重匱乏；（3）甲基化、羥基化、糖基化及酰基化等催化作用對花青素的形成和運(yùn)輸有重要作用，但光信號是否影響這些修飾過程尚不清楚；（4）園藝植物果實中花青素的積累通常受光照、溫度等環(huán)境因素共同作用的影響，探究光環(huán)境與其他環(huán)境因子在調(diào)控花青素中的互作機(jī)制，對合理利用設(shè)施環(huán)境調(diào)控手段，精準(zhǔn)調(diào)控園藝植物花青素的形成，進(jìn)而改良觀賞植物花色、提高園藝產(chǎn)品的營養(yǎng)價值有重要的指導(dǎo)意義。

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Light Regulation of Anthocyanin Biosynthesis in Horticultural Crops

WANG Feng, WANG XiuJie, ZHAO ShengNan, YAN JiaRong, BU Xin, ZHANG Ying, LIU YuFeng, XU Tao, QI MingFang, QI HongYan, LI TianLai

(College of Horticulture, Shenyang Agricultural University/The State Education Ministry and Liaoning Provincial Key Laboratory of Protected Horticulture/National & Local Joint Engineering Research Center of Northern Horticultural Facilities Design & Application Technology (Liaoning), Shenyang 110866)

Anthocyanins are among the most important flavonoid compounds in plants, which play significant roles in color formation of plant organ, such as flower and fruits, as well as antioxidant process. Light is one of the most important environmental factors affecting anthocyanin biosynthesis pathway, but it still remains unclear in the mechanism and signaling networks of light regulation of anthocyanin. This review briefly introduced the anthocyanin biosynthesis and transportation pathway, and summarized the molecular mechanism of anthocyanin transcriptional regulation by three kinds of transcription factors, including MYB, bHLH and WDR. In addition, it emphasized on the light signaling regulation of anthocyanin biosynthesis. The researches showed that the light environment (light intensity, light quality, and light duration) regulated the biosynthetic process of anthocyanin mainly through different light receptors (UVR8, CRYs, PHOTs, and PHYs), which affected the ubiquitination ability of COP1, the stability of HY5, and the stability of other light signal transcription factors, such as the phytochrome-interacting factors (PIFs). On the one hand, these light signal factors directly could bind to the promoters of,and, activate or inhibit these genes expression and then regulate the synthesis of anthocyanin. On the other hand, these light signal factors interacted with proteins of MYB, bHLH and WDR, affecting the stability of the MBW complex formed by them. In addition, these light signaling factors could also regulate anthocyanin synthesis through MBW independent pathways, such as HY5 also affect anthocyanin biosynthesis by regulating. In addition, some unknown light signaling factors might directly or indirectly regulate anthocyanin synthesis genes and interacting with some vacuolar membranes proteins in a MBW independent manner, to change vacuolar acidification and regulate anthocyanin synthesis. At the same time, light signaling factors also affected some factors in the photosynthetic electron transport chain through MBW dependent or MBW independent pathways, then affected anthocyanin synthesis in plants. How these pathways were coordinated and which pathway was preferentially responded by light environments (light intensity, light quality, light duration)? This paper provided a basis to further investigate the molecular mechanism regulating anthocyanin biosynthesis by light signalings. The study explored the effective ways and target molecules for light regulation of anthocyanin accumulation, andcreated opportunities for the development of anthocyanin-rich horticultural crops through genetic and metabolic engineering, and light environmental management.

light; anthocyanins; transcription factor; transcriptional regulation; horticultural crops

10.3864/j.issn.0578-1752.2020.23.015

2020-04-06；

2020-06-30

國家自然科學(xué)基金（31801904）、遼寧省“興遼英才計劃”（XLYC1807020）、遼寧省高等學(xué)校創(chuàng)新人才支持計劃（LR2018027）、遼寧省博士啟動基金（20180540094）、沈陽市中青年科技創(chuàng)新人才支持計劃（RC200449）、國家重點研發(fā)計劃（2018YFD1000800，2019YFD1000300）、國家現(xiàn)代農(nóng)業(yè)產(chǎn)業(yè)技術(shù)體系建設(shè)專項（CARS-23-C01）、遼寧省“百千萬人才工程”（LNBQW2018W0483）、沈陽農(nóng)業(yè)大學(xué)科研啟動基金（880418039）

通信作者王峰，E-mail：fengwang@syau.edu.cn。通信作者李天來，E-mail：tianlaili@126.com

（責(zé)任編輯 趙伶俐）