植物学报 ›› 2016, Vol. 51 ›› Issue (1): 107-119.DOI: 10.11983/CBB15059
祝志欣1,2, 鲁迎青1,*(
)
首页收稿日期:2015-03-26
接受日期:2015-07-17
出版日期:2016-01-01
发布日期:2016-02-01
通讯作者:
鲁迎青
作者简介:? 共同第一作者
基金资助:Zhixin Zhu1,2, Yingqing Lu1*
Received:2015-03-26
Accepted:2015-07-17
Online:2016-01-01
Published:2016-02-01
Contact:
Lu Yingqing
About author:? These authors contributed equally to this paper
摘要： 花青素是种子植物呈色的重要色素, 由一系列结构基因编码的酶(CHS、CHI、F3H、F3'H、F3'5'H、DFR、ANS和3GT)催化而成, 随后经过各种修饰被转运至液泡等部位储存。各类器官中差异表达的MYB、bHLH和WDR三种调控因子通过形成MBW复合体直接正调控以上结构基因的表达。这个过程涉及的基因变异常会导致植物的各种颜色变异。在生活中人们广泛利用这些变异品种, 取其丰富色味。造成颜色变异的具体分子机制在很多情况下还不清楚, 但日益积累的个例研究为其中的规律性提供了基础数据。该文概述了花青素的合成、转运过程及其转录调控机制, 探讨了研究颜色变异品种的常用思路及方法。在总结近年工作的基础上, 对生活中常见蔬菜、水果和花卉的颜色变异品种的分子机制进行了综述。
祝志欣, 鲁迎青. 花青素代谢途径与植物颜色变异. 植物学报, 2016, 51(1): 107-119.
Zhixin Zhu, Yingqing Lu. Plant Color Mutants and the Anthocyanin Pathway. Chinese Bulletin of Botany, 2016, 51(1): 107-119.
图1 类黄酮代谢途径中花青素的合成示意图 花青素合成途径的中间底物也可被导向类黄酮途径的其它分支(灰色箭头所示)。
Figure 1 The flavonoid biosynthetic pathway leading to the production of anthocyaninThe intermediate substrates can be canalized to other branches of the flavonoid pathway, which were indicated with grey arrows.
| 物种 | 花青素相关MBW复合物的成员 | ||
|---|---|---|---|
| MYB | bHLH | WDR | |
| 玉米(Zea mays) | ZmC1, ZmPl | ZmR, ZmB | ZmPAC1 |
| 拟南芥(Arabidopsis thaliana) | AtPAP1, AtPAP2 | AtGL3, AtEGL3 | AtTTG1 |
| 矮牵牛(Petunia hybrida ) | PhAN2, PhAN4 | PhAN1 | PhAN11 |
| 圆叶牵牛(Ipomoea purpurea) | IpMYB1 | IpbHLH2 | IpWDR1 |
表1 常见模式研究物种中花青素相关MBW复合物的成员
Table 1 Members of the MBW complex in anthocyanin model species
| 物种 | 花青素相关MBW复合物的成员 | ||
|---|---|---|---|
| MYB | bHLH | WDR | |
| 玉米(Zea mays) | ZmC1, ZmPl | ZmR, ZmB | ZmPAC1 |
| 拟南芥(Arabidopsis thaliana) | AtPAP1, AtPAP2 | AtGL3, AtEGL3 | AtTTG1 |
| 矮牵牛(Petunia hybrida ) | PhAN2, PhAN4 | PhAN1 | PhAN11 |
| 圆叶牵牛(Ipomoea purpurea) | IpMYB1 | IpbHLH2 | IpWDR1 |
图2 花青素代谢途径转录激活假说示意图 花青素相关MYB被诱导表达后, 可反馈性激活bHLH和WDR的表达, 并组成MBW复合物。MBW复合物的激活能力显著高于MYB本身, 其靶基因包括花青素合成途径的整套基因和之后的部分修饰与转运相关基因。MBW复合物对应2个识别序列, 分别为MRE (MYB识别序列)和BRE (bHLH识别序列), 它们分别遵循ANCNN(C/A)C和CACN(A/C/T)(G/T)序列模式(Zhu et al., 2015)。不论是MBW复合物功能异常, 还是靶基因启动子上MRE和BRE序列突变, 都可能导致花青素的合成异常。
Figure 2 The hypothesized regulatory mechanism for the activation of the anthocyanin pathwayOnce the anthocyanin MYB was induced by signals, it may activate the expression of bHLH and WDR, forming the MBW complex all together. The MBW complex has much higher activation ability than MYB alone. The targets of the MBW complex include the whole set of the core structural genes of the anthocyanin pathway, and likely also comprising genes encoding relevant modification and transportation enzymes. There were two classes of cis elements for the MBW complex, the MRE (MYB-recognizing elements) and BRE (bHLH-recognizing elements), which were in the form of ANCNN(C/A)C and CACN(A/C/T)(G/T) respectively (Zhu et al., 2015). Either the functional disorder of the MBW complex or the mutations at MRE or/and BRE of the target promoters may cause abnormal anthocyanin biosynthesis.
图3 颜色变异品种的常见研究思路及方法
Figure 3 Common strategies for dissecting the molecular basis of a plant color mutant
图4 生活中与花青素代谢途径相关的常见颜色变异品种(部分图改自相应参考文献) (A) 水稻红米(Furukawa et al., 2007); (B) 杂色玉米(Lazarow et al., 2013); (C) 紫皮、绿皮葡萄; (D) 紫薯(Mano et al., 2007); (E) 各色圆叶牵牛; (F) 各色大丽花(Ohno et al., 2011); (G) 红肉苹果(Espley et al., 2009); (H) 彩叶草(Nguyen and Cin, 2009); (I) 紫色卷心菜(Yuan et al., 2009); (J) 紫色花椰菜(Chiu et al., 2010); (K) 红斑文心兰
Figure 4 Examples of commonly seen plant color mutants (some graphs were modified from relative references) (A) Red rice (Furukawa et al., 2007); (B) Variegated corn (Lazarow et al., 2013); (C) White grape; (D) Purple sweet potato (Mano et al., 2007); (E) Common morning glory; (F) Dahlia (Ohno et al., 2011); (G) Red fleshed apple (Espley et al., 2009); (H) Coleus (Nguyen and Cin, 2009); (I) Purple cabbage (Yuan et al., 2009); (J) Purple cauliflower (Chiu et al., 2010); (K) Red spotted dancing-doll orchid
| 植物 类别 | 物种名 | 颜色变异品种 | 突变位点 | 相关分子机制 | 参考文献 |
|---|---|---|---|---|---|
| 粮食 作物 | 水稻 (Oryza sativa) | 红米 ( | bHLH类调控基因; 结构基因DFR | 红色稻米含原花青素, 推测为祖先型性状。白色稻米含有Rc或Rd位点的突变。其中Rc为bHLH类转录因子, 而Rd属于结构基因DFR。 | Sweeney et al., 2006; Furukawa et al., 2007 |
| 玉米 (Zea mays) | 红色及杂色品种 ( | MYB或bHLH类调控基因; 众多结构基因 | 玉米粒的颜色由表皮的鞣红和糊粉层的花青素共同构成。其祖先状态被认为是红紫色品种。玉米基因组中转座子在类黄酮相关的基因附近频繁活动, 使得这些基因常常失去或恢复活性, 而形成各种杂色表型。 | McClintock, 1950; Mol et al., 1998; Lazarow et al., 2013 | |
| 红薯 (Ipomoea batatas) | 紫薯 ( | MYB类调控基因 | 日常食用的红薯块根为白色到橙色, 不含色素或含类胡萝卜素。紫薯中则积累了不同程度的花青素。研究表明, 这是由于IbMYB1在紫薯块根中特异表达, 使块根中的花青素代谢途径被上调所致。 | Mano et al., 2007 | |
| 蔬菜 | 花椰菜 (Brassica oleracea var. botrytis) | 紫色花 椰菜 ( | MYB类调控基因 | 紫色品种中BoMYB2的启动子被插入1个转座子, 引起BoMYB2在球茎中大量表达, 球茎中的花青素代谢途径被上调。 | Chiu et al., 2010 |
| 卷心菜 (Brassica oleracea var. capitata) | 紫色卷 心菜 ( | MYB或bHLH类调控基因 | 紫色卷心菜中花青素代谢途径的正调控因子BoMYB2和 BoTT8表达显著提高, 而负调控因子BoMYB3的表达反而有所降低。 | Yuan et al., 2009 | |
| 水果 | 苹果 (Malus x domestica) | 红肉苹果 ( | MYB类调控基因 | 苹果一般为红皮白肉, 但自然界中也有红肉的品种。研究表明, 红肉苹果中MdMYB10的启动子近端区一处短片段发生了串联重复, 而此短片段中正好含有Md- MYB10蛋白的自身调控元件, 造成了MdMYB10基因可以正调控自身的表达, 形成无节制的自身正反馈。 | Espley et al., 2009; Wang et al., 2013a |
| 葡萄 (Vitis vinifera) | 紫皮和绿皮葡萄 ( | MYB类调控基因 | 葡萄总体上分为红黑皮和绿皮两个类群, 葡萄皮的颜色直接决定了葡萄酒的颜色。研究表明, 红黑皮为葡萄的祖先状态, 而白皮品种是由于其花青素特异调控基因Vv- MYBA1和VvMYBA2同时突变导致花青素特异结构基因3GT不能正常表达引起。 | Walker et al., 2007 | |
| 观赏 植物 | 文心兰 (Oncidium ‘Gower Ramsey’) | 花瓣上的花纹 ( | MYB类调控基因 | 文心兰的花为黄色背景带红色纹理。研究表明, 黄色背景部位缺乏花青素, 是由于OgMYB1在黄色部分不表达, 导致花青素代谢途径的2个结构基因OgCHI和Og- DFR不能表达。 | Chiou and Yeh, 2008 |
| 圆叶牵牛 (Ipomoea purpurea) | 多种花色着色形式 ( | MYB、bHLH或WDR类调控基因; 众多结构基因 | 圆叶牵牛具有极其多样的着色模式, 其变异涉及花青素代谢途径几乎所有的调控基因和结构基因, 大都由转座子的活动引起。它的野生型为蓝紫色, 红色为F3'H的突变引起, 白色和杂色品种则可以由众多其它基因上的突变引起。 | Hoshino et al., 2001, 2003; Clegg and Durbin, 2003; Morita et al., 2006; Park et al., 2007 | |
| 大丽花 (Dahlia variabilis) | 多种花色着色形式 ( | bHLH类调控基因 | 野生型大丽花为橙红色, 但其bHLH类调控基因被插入1个转座子后就可以形成黄底橙斑的杂色形式。这种杂色大丽花由于转座子的切离会发生后代花色分离。转座子非移码性切离可以产生完全回复的橙红色表型, 移码性切离会产生完全突变的黄色表型; 转座子不切离则后代将保留亲代黄底橙斑的杂色形式。 | Ohno et al., 2011 | |
| 彩叶草 (Solenostemon scutellarioides) | 叶片具丰富的着色形式 ( | 发育和环境信号联合导致花青素的不同程度表达引起 | 彩叶草由于叶色绚丽也被列为观赏植物。其多样的叶色其实是叶绿素和花青素不同比例组合的结果, 这种比例组合受到发育和环境信号的共同调控。 | Nguyen and Cin, 2009 |
表2 生活中常见颜色变异品种的已知分子机制
Table 2 Known molecular mechanisms responsible for common color mutants in cultivated plants
| 植物 类别 | 物种名 | 颜色变异品种 | 突变位点 | 相关分子机制 | 参考文献 |
|---|---|---|---|---|---|
| 粮食 作物 | 水稻 (Oryza sativa) | 红米 ( | bHLH类调控基因; 结构基因DFR | 红色稻米含原花青素, 推测为祖先型性状。白色稻米含有Rc或Rd位点的突变。其中Rc为bHLH类转录因子, 而Rd属于结构基因DFR。 | Sweeney et al., 2006; Furukawa et al., 2007 |
| 玉米 (Zea mays) | 红色及杂色品种 ( | MYB或bHLH类调控基因; 众多结构基因 | 玉米粒的颜色由表皮的鞣红和糊粉层的花青素共同构成。其祖先状态被认为是红紫色品种。玉米基因组中转座子在类黄酮相关的基因附近频繁活动, 使得这些基因常常失去或恢复活性, 而形成各种杂色表型。 | McClintock, 1950; Mol et al., 1998; Lazarow et al., 2013 | |
| 红薯 (Ipomoea batatas) | 紫薯 ( | MYB类调控基因 | 日常食用的红薯块根为白色到橙色, 不含色素或含类胡萝卜素。紫薯中则积累了不同程度的花青素。研究表明, 这是由于IbMYB1在紫薯块根中特异表达, 使块根中的花青素代谢途径被上调所致。 | Mano et al., 2007 | |
| 蔬菜 | 花椰菜 (Brassica oleracea var. botrytis) | 紫色花 椰菜 ( | MYB类调控基因 | 紫色品种中BoMYB2的启动子被插入1个转座子, 引起BoMYB2在球茎中大量表达, 球茎中的花青素代谢途径被上调。 | Chiu et al., 2010 |
| 卷心菜 (Brassica oleracea var. capitata) | 紫色卷 心菜 ( | MYB或bHLH类调控基因 | 紫色卷心菜中花青素代谢途径的正调控因子BoMYB2和 BoTT8表达显著提高, 而负调控因子BoMYB3的表达反而有所降低。 | Yuan et al., 2009 | |
| 水果 | 苹果 (Malus x domestica) | 红肉苹果 ( | MYB类调控基因 | 苹果一般为红皮白肉, 但自然界中也有红肉的品种。研究表明, 红肉苹果中MdMYB10的启动子近端区一处短片段发生了串联重复, 而此短片段中正好含有Md- MYB10蛋白的自身调控元件, 造成了MdMYB10基因可以正调控自身的表达, 形成无节制的自身正反馈。 | Espley et al., 2009; Wang et al., 2013a |
| 葡萄 (Vitis vinifera) | 紫皮和绿皮葡萄 ( | MYB类调控基因 | 葡萄总体上分为红黑皮和绿皮两个类群, 葡萄皮的颜色直接决定了葡萄酒的颜色。研究表明, 红黑皮为葡萄的祖先状态, 而白皮品种是由于其花青素特异调控基因Vv- MYBA1和VvMYBA2同时突变导致花青素特异结构基因3GT不能正常表达引起。 | Walker et al., 2007 | |
| 观赏 植物 | 文心兰 (Oncidium ‘Gower Ramsey’) | 花瓣上的花纹 ( | MYB类调控基因 | 文心兰的花为黄色背景带红色纹理。研究表明, 黄色背景部位缺乏花青素, 是由于OgMYB1在黄色部分不表达, 导致花青素代谢途径的2个结构基因OgCHI和Og- DFR不能表达。 | Chiou and Yeh, 2008 |
| 圆叶牵牛 (Ipomoea purpurea) | 多种花色着色形式 ( | MYB、bHLH或WDR类调控基因; 众多结构基因 | 圆叶牵牛具有极其多样的着色模式, 其变异涉及花青素代谢途径几乎所有的调控基因和结构基因, 大都由转座子的活动引起。它的野生型为蓝紫色, 红色为F3'H的突变引起, 白色和杂色品种则可以由众多其它基因上的突变引起。 | Hoshino et al., 2001, 2003; Clegg and Durbin, 2003; Morita et al., 2006; Park et al., 2007 | |
| 大丽花 (Dahlia variabilis) | 多种花色着色形式 ( | bHLH类调控基因 | 野生型大丽花为橙红色, 但其bHLH类调控基因被插入1个转座子后就可以形成黄底橙斑的杂色形式。这种杂色大丽花由于转座子的切离会发生后代花色分离。转座子非移码性切离可以产生完全回复的橙红色表型, 移码性切离会产生完全突变的黄色表型; 转座子不切离则后代将保留亲代黄底橙斑的杂色形式。 | Ohno et al., 2011 | |
| 彩叶草 (Solenostemon scutellarioides) | 叶片具丰富的着色形式 ( | 发育和环境信号联合导致花青素的不同程度表达引起 | 彩叶草由于叶色绚丽也被列为观赏植物。其多样的叶色其实是叶绿素和花青素不同比例组合的结果, 这种比例组合受到发育和环境信号的共同调控。 | Nguyen and Cin, 2009 |
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