园艺学报 ›› 2021, Vol. 48 ›› Issue (10): 2057-2072.doi: 10.16420/j.issn.0513-353x.2021-0444
郑清冬, 王艺, 欧悦, 柯玉洁, 姚亚合, 王梦洁, 陈嘉忆, 艾叶*()
收稿日期:
2021-06-23
修回日期:
2021-08-26
出版日期:
2021-10-25
发布日期:
2021-11-01
通讯作者:
艾叶
E-mail:aiyefafu@163.com
基金资助:
ZHENG Qingdong, WANG Yi, OU Yue, KE Yujie, YAO Yahe, WANG Mengjie, CHEN Jiayi, AI Ye*()
Received:
2021-06-23
Revised:
2021-08-26
Online:
2021-10-25
Published:
2021-11-01
Contact:
AI Ye
E-mail:aiyefafu@163.com
摘要:
对兰科植物的花色素种类、花青素苷和类胡萝卜素合成途径相关基因的研究进展进行了综述,并对今后的研究方向进行展望,以期为兰科植物花色相关分子机制的研究及花色品种培育提供参考信息。
中图分类号:
郑清冬, 王艺, 欧悦, 柯玉洁, 姚亚合, 王梦洁, 陈嘉忆, 艾叶. 兰科植物花色相关基因研究进展[J]. 园艺学报, 2021, 48(10): 2057-2072.
ZHENG Qingdong, WANG Yi, OU Yue, KE Yujie, YAO Yahe, WANG Mengjie, CHEN Jiayi, AI Ye. Research Advances of Genes Responsible for Flower Colors in Orchidaceae[J]. Acta Horticulturae Sinica, 2021, 48(10): 2057-2072.
图1 花青素苷合成途径 PAL:苯丙氨酸解氨酶;C4H:肉桂酸4-羟基化酶;4CL:4-香豆酸辅酶A连接酶;CHS:查尔酮合成酶;CHI:查尔酮异构酶;F3H:黄烷酮-3-羟化酶;F3′5′H:类黄酮-3′,5′-羟基化酶;F3′H:类黄酮-3′-羟化酶;FLS:黄酮醇合成酶;DFR:二氢黄酮醇还原酶;ANS:花青素合成酶;LAR:无色花青素还原酶;ANR:花青素还原酶;UFGT:类黄酮糖基转移酶;OMT:O-甲基转移酶。
Fig. 1 Biosynthesis pathway of anthocyanins PAL:Phenylalanin ammonialyase;C4H:Cinnamate 4-hydroxylase;4CL:4-coumarate;coenzyme ligase;CHS:Chalcone synthase;CHI:Chalcone synthase;F3H:Flavanone-3-hydroxylase;F3′5′H:Dihydroflavonoid-3′,5′-hydroxylase;F3'H:Dihydroflavonoid-3′-hydroxylase;FLS:Flavonol synthase;DFR:Dihydroflavonol reductase;ANS:Anthocyanin synthase;LAR:Leucoanthocyanidin reductase;ANR:Anthocyanidin reductase;UFGT:UDP-glucose flavonoid3-glucosyltransferase;OMT:O-methyltransferases.
类型 Type | 基因名称 Gene name | 来源植物 Original plant | 基因功能 Gene function | 参考文献 Reference |
---|---|---|---|---|
结构基因 Structural gene | CHS | 铁皮石斛 Dendrobium officinale | 参与类黄酮的合成 Participate in the synthesis of flavonoids | 孟衡玲 等, |
OgCHS | 文心兰 Oncidium‘Honey Dollp’ | 瞬时表达产生花青素苷 Transient expression to produce anthocyanins | Liu et al., | |
PcCHS | 兜兰 Paphiopedilum concolor Pfitz | 对花瓣中的紫色斑点起着调控作用 It can regulate the purple spots in petals | 李冬梅 等, | |
CHS | 建兰‘龙福星蝶’Cymbidium ensifolium‘Longfuxingdie’ | 在红色区域表达,在黄绿色区域不表达 It is expressed in red area,but not in yellow green area | 李文建 等, | |
OgCHI | 文心兰 Oncidium Gower Ramsey | 使黄色唇瓣产生花青素苷 It makes the yellow lips produce anthocyanins | Chiou & Yeh, | |
CHI2 | 杂交石斛兰 Dendrobium Sirin classic | 低表达导致颜色强度很低 Low expression leads to faded color | Kriangphan et al., | |
Va-F3H1 | 万代兰 Vanda hybrid | 在白色品种中表达量非常低,在淡紫色品种中表达量较高 The expression is very low in white varieties,but high in light purple varieties | Junka et al., | |
F3H | 石斛兰 Dendrobium officinale | 其表达影响二氢黄酮醇含量,进而影响花青素苷的积累 The expression affected the content of dihydroflavonol,and then affect the accumulation of anthocyanins | Zhan et al., | |
DmF3′5′H | 念珠石斛 Dendrobium moniliforme | 使白色花被片产生了紫红色斑点 The white perianth produces purplish red spots | Whang et al., | |
F3′5′H | 蝴蝶兰 Phalaenopsis hybrid | 转基因植株花瓣颜色由粉红色变为红紫色 The petal color of transgenic plants changed from pink to red purple | Su & Hsu, | |
PhF3′H | 蝴蝶兰 Phalaenopsis | 在红色蝴蝶兰中的表达量远高于黄色蝴蝶兰 The expression level in red Phalaenopsis is much higher than that in yellow Phalaenopsis | 杨玉霞 等, | |
PsDFR | 蝴蝶兰 Phalaenopsis‘Everspring Fairy’ | 低表达导致花瓣和萼片ABP受阻,没有花色素合成 Low expression of ABP in petals and sepals resulted in inhibition of anthocyanin synthesis | Ma et al., | |
PeUFGT3 | 蝴蝶兰 Phalaenopsis | 在红色花色形成过程中起重要作用 It plays an important role in the formation of red flower | Chen et al., | |
CHS,DFR, ANS | 大花蕙兰 Cymbidium hybrida | 粉红色品种中的表达高于黄色、绿色品种 The expression in pink was higher than that in yellow and green | Wang et al., | |
ANS | 万代兰 Vanda hybrid | 白色的形成可能与其缺失有关 The formation of white may be related to the absence of ANS | Junka et al., | |
调控基因Regulatory gene | OgMYB1 | 文心兰 Oncidium Gower Ramsey | 激活OgCHI和OgDFR转录诱导红色色素的形成 Activation of OgCHI and OgDFR transcription induces red pigment formation | Chiou & Yeh, |
MYB | 蝴蝶兰 Phalaenopsis‘Everspring Fairy’ | 仅在紫色区域具有高表达,在白色区域没有表达 It is highly express only in the purple region,but not in the white region | Ma et al., | |
类型 Type | 基因名称 Gene name | 来源植物 Original plant | 基因功能 Gene function | 参考文献 Reference |
PeMYB2, PeMYB11, PeMYB12 | 小兰屿蝴蝶兰 Phalaenopsis equestris | 控制萼片和花瓣的底色、斑点和网纹 The color patterns of sepals and petals were controlled by the background,spots and reticulation | Hsu et al., | |
PeMYB11 | 蝴蝶兰 Phalaenopsis Yushan Little Pearl | 高表达产生紫黑色斑点 Purple black spots were produced by overexpression of PeMYB11 | Hsu et al., | |
PeMYB7 PeMYB11 | 蝴蝶兰 Phalaenopsis‘Panda’ | 正向调控PeF3H、PeF3'H和PeANS的表达 It promotes the expression of PeF3H,PeF3′H and PeANS | Zhao et al., | |
RcPAP1 RcPAP2 | 卡特兰 Cattleya hybrid‘KOVA’ | 使蝴蝶兰白色花被片变成了紫红色 It turns white perianth of Phalaenopsis to purplish red | Li et al., | |
MYBx1 | 蝴蝶兰 Phalaenopsis‘Big Chili.’ | 编码R3-MYB蛋白,抑制花青素苷的积累 It encodes R3-MYB protein and inhibits anthocyanin accumulation | Fu et al., | |
MYB10 | 四川独蒜兰 Pleione limprichtii | 调节PlFLS的表达,作为抑制花青素苷形成的阻遏物 It can regulate the expression of PlFLS as a repressor of anthocyanin formation | Zhang et al., | |
MYB10 | 蝴蝶石斛兰 Dendrobium phalaenopsis | 负调控DFR、ANS、F3H、F3′H的表达 It inhibits the expression of DFR,ANS,F3H and F3′H | 丁灵,2016 | |
CyMYB1 CybHLH1 CybHLH2 | 兰属 Cymbidium‘Mystique’ | 形成复合物可以激活CyDFR和CyANS The formation of complexes can activate CyDFR and CyANS | Nakatsuka et al., | |
DhMYB2 DhbHLH1 | 石斛杂交种 Dendrobium hybrids orchid | 调节DhF3H、DhDFR和DhANS,导致白色花瓣产生紫色斑点 Regulation of DhF3H,DhDFR and DhANS genes resulted in purple spots on white petals | Li et al., | |
bHLH, MYB | 大花蕙兰 Cymbidium hybrid‘Jung Frau dos Pueblos’ | 同时过表达可产生大量花青素苷 Overexpression of these two genes at the same time can produce a large amount of anthocyanins | Albert et al., | |
MYB4 bHLH1 | 蝴蝶石斛兰 Dendrobium phalaenopsis | bHLH1与MYB4形成互作的关系调控花青素苷生成 The interaction between bHLH1 and MYB4 formation regulates anthocyanin production | 丁灵,2016 |
表1 兰科植物花青素苷合成相关基因及其功能
Table 1 Anthocyanin biosynthesis related genes and their functions in orchids
类型 Type | 基因名称 Gene name | 来源植物 Original plant | 基因功能 Gene function | 参考文献 Reference |
---|---|---|---|---|
结构基因 Structural gene | CHS | 铁皮石斛 Dendrobium officinale | 参与类黄酮的合成 Participate in the synthesis of flavonoids | 孟衡玲 等, |
OgCHS | 文心兰 Oncidium‘Honey Dollp’ | 瞬时表达产生花青素苷 Transient expression to produce anthocyanins | Liu et al., | |
PcCHS | 兜兰 Paphiopedilum concolor Pfitz | 对花瓣中的紫色斑点起着调控作用 It can regulate the purple spots in petals | 李冬梅 等, | |
CHS | 建兰‘龙福星蝶’Cymbidium ensifolium‘Longfuxingdie’ | 在红色区域表达,在黄绿色区域不表达 It is expressed in red area,but not in yellow green area | 李文建 等, | |
OgCHI | 文心兰 Oncidium Gower Ramsey | 使黄色唇瓣产生花青素苷 It makes the yellow lips produce anthocyanins | Chiou & Yeh, | |
CHI2 | 杂交石斛兰 Dendrobium Sirin classic | 低表达导致颜色强度很低 Low expression leads to faded color | Kriangphan et al., | |
Va-F3H1 | 万代兰 Vanda hybrid | 在白色品种中表达量非常低,在淡紫色品种中表达量较高 The expression is very low in white varieties,but high in light purple varieties | Junka et al., | |
F3H | 石斛兰 Dendrobium officinale | 其表达影响二氢黄酮醇含量,进而影响花青素苷的积累 The expression affected the content of dihydroflavonol,and then affect the accumulation of anthocyanins | Zhan et al., | |
DmF3′5′H | 念珠石斛 Dendrobium moniliforme | 使白色花被片产生了紫红色斑点 The white perianth produces purplish red spots | Whang et al., | |
F3′5′H | 蝴蝶兰 Phalaenopsis hybrid | 转基因植株花瓣颜色由粉红色变为红紫色 The petal color of transgenic plants changed from pink to red purple | Su & Hsu, | |
PhF3′H | 蝴蝶兰 Phalaenopsis | 在红色蝴蝶兰中的表达量远高于黄色蝴蝶兰 The expression level in red Phalaenopsis is much higher than that in yellow Phalaenopsis | 杨玉霞 等, | |
PsDFR | 蝴蝶兰 Phalaenopsis‘Everspring Fairy’ | 低表达导致花瓣和萼片ABP受阻,没有花色素合成 Low expression of ABP in petals and sepals resulted in inhibition of anthocyanin synthesis | Ma et al., | |
PeUFGT3 | 蝴蝶兰 Phalaenopsis | 在红色花色形成过程中起重要作用 It plays an important role in the formation of red flower | Chen et al., | |
CHS,DFR, ANS | 大花蕙兰 Cymbidium hybrida | 粉红色品种中的表达高于黄色、绿色品种 The expression in pink was higher than that in yellow and green | Wang et al., | |
ANS | 万代兰 Vanda hybrid | 白色的形成可能与其缺失有关 The formation of white may be related to the absence of ANS | Junka et al., | |
调控基因Regulatory gene | OgMYB1 | 文心兰 Oncidium Gower Ramsey | 激活OgCHI和OgDFR转录诱导红色色素的形成 Activation of OgCHI and OgDFR transcription induces red pigment formation | Chiou & Yeh, |
MYB | 蝴蝶兰 Phalaenopsis‘Everspring Fairy’ | 仅在紫色区域具有高表达,在白色区域没有表达 It is highly express only in the purple region,but not in the white region | Ma et al., | |
类型 Type | 基因名称 Gene name | 来源植物 Original plant | 基因功能 Gene function | 参考文献 Reference |
PeMYB2, PeMYB11, PeMYB12 | 小兰屿蝴蝶兰 Phalaenopsis equestris | 控制萼片和花瓣的底色、斑点和网纹 The color patterns of sepals and petals were controlled by the background,spots and reticulation | Hsu et al., | |
PeMYB11 | 蝴蝶兰 Phalaenopsis Yushan Little Pearl | 高表达产生紫黑色斑点 Purple black spots were produced by overexpression of PeMYB11 | Hsu et al., | |
PeMYB7 PeMYB11 | 蝴蝶兰 Phalaenopsis‘Panda’ | 正向调控PeF3H、PeF3'H和PeANS的表达 It promotes the expression of PeF3H,PeF3′H and PeANS | Zhao et al., | |
RcPAP1 RcPAP2 | 卡特兰 Cattleya hybrid‘KOVA’ | 使蝴蝶兰白色花被片变成了紫红色 It turns white perianth of Phalaenopsis to purplish red | Li et al., | |
MYBx1 | 蝴蝶兰 Phalaenopsis‘Big Chili.’ | 编码R3-MYB蛋白,抑制花青素苷的积累 It encodes R3-MYB protein and inhibits anthocyanin accumulation | Fu et al., | |
MYB10 | 四川独蒜兰 Pleione limprichtii | 调节PlFLS的表达,作为抑制花青素苷形成的阻遏物 It can regulate the expression of PlFLS as a repressor of anthocyanin formation | Zhang et al., | |
MYB10 | 蝴蝶石斛兰 Dendrobium phalaenopsis | 负调控DFR、ANS、F3H、F3′H的表达 It inhibits the expression of DFR,ANS,F3H and F3′H | 丁灵,2016 | |
CyMYB1 CybHLH1 CybHLH2 | 兰属 Cymbidium‘Mystique’ | 形成复合物可以激活CyDFR和CyANS The formation of complexes can activate CyDFR and CyANS | Nakatsuka et al., | |
DhMYB2 DhbHLH1 | 石斛杂交种 Dendrobium hybrids orchid | 调节DhF3H、DhDFR和DhANS,导致白色花瓣产生紫色斑点 Regulation of DhF3H,DhDFR and DhANS genes resulted in purple spots on white petals | Li et al., | |
bHLH, MYB | 大花蕙兰 Cymbidium hybrid‘Jung Frau dos Pueblos’ | 同时过表达可产生大量花青素苷 Overexpression of these two genes at the same time can produce a large amount of anthocyanins | Albert et al., | |
MYB4 bHLH1 | 蝴蝶石斛兰 Dendrobium phalaenopsis | bHLH1与MYB4形成互作的关系调控花青素苷生成 The interaction between bHLH1 and MYB4 formation regulates anthocyanin production | 丁灵,2016 |
图2 类胡萝卜素生物合成途径 PSY:八氢番茄红素合成酶;PDS:八氢番茄红素脱氢酶;ZDS:ζ-胡萝卜素脱氢酶;CRTISO:类胡萝卜素异构酶;LYCE:番茄红素ε-环化酶;LYCB:番茄红素β-环化酶;BCH:β-胡萝卜素羟化酶;VDE:紫黄质脱环氧化酶;ZEP:玉米黄质环氧酶;CCS:辣椒红素合酶;NSY:新黄质合成酶。
Fig. 2 Carotenoid biosynthesis pathway PSY:Phytoene synthase;PDS:Phytoene desaturase;ZDS:ζ-carotene desaturase;CRTISO:Carotenoid isomerase;LYCE:Lycopene ε-cyclase;LYCB:Lycopene β-cyclase;BCH:β-carotene hydroxylase,VDE:Violaxanthinde-epoxidase;ZEP:Zeaxanthin epoxidase;CCS:Capsanthin/Capsorubin synthase;NSY:Neoxanthin synthase.
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