浙江农业学报 ›› 2021, Vol. 33 ›› Issue (6): 1149-1158.DOI: 10.3969/j.issn.1004-1524.2021.06.21
• 综述 • 上一篇
MADS-box基因家族调控植物花器官发育研究进展
王莹(
), 穆艳霞, 王锦*()
- 西南林业大学 园林园艺学院,云南 昆明 650224
-
收稿日期:2020-09-16出版日期:2021-06-25发布日期:2021-06-25 -
通讯作者:王锦 -
作者简介:*王锦,E-mail: 908505685@qq.com
王莹(1986—),女,河南新乡人,博士研究生,讲师,主要从事园林植物与观赏园艺植物方面的研究。E-mail: 327281514@qq.com -
基金资助:国家林业局林业科技成果国家级推广项目(201528);云南省教育厅科学研究基金(2021Y253)
Research progress of floral development regulation by MADS-box gene family
WANG Ying(), MU Yanxia, WANG Jin*()
- College of Landscape Architecture and Horticulture Science, Southwest Forestry University, Kunming 650224, China
-
Received:2020-09-16Online:2021-06-25Published:2021-06-25 -
Contact:WANG Jin
摘要：
花器官的发生发育与形态构造是一个高度复杂的项目,涉及众多因素及其相互作用,MADS-box基因作为其中的关键转录因子可以改变整个发育过程,因而成为花器官研究最广泛的基因家族。通过梳理目前MADS-box基因对花器官发生、分化和形态建成方面的调控作用,为了解花发育程序和该基因家族的一般转录调控机制提供新见解,也为进一步深入挖掘该家族基因和完善花发育调控理论提供参考。
中图分类号:
引用本文
王莹, 穆艳霞, 王锦. MADS-box基因家族调控植物花器官发育研究进展[J]. 浙江农业学报, 2021, 33(6): 1149-1158.
WANG Ying, MU Yanxia, WANG Jin. Research progress of floral development regulation by MADS-box gene family[J]. Acta Agriculturae Zhejiangensis, 2021, 33(6): 1149-1158.
图1 MADS-box 蛋白分类与结构[7,8,13-14]
Fig.1 Classification and structure of MADS-box proteins
图1 MADS-box 蛋白分类与结构[7,8,13-14]
Fig.1 Classification and structure of MADS-box proteins
表1 MADS-box基因及其功能[2,14,16-18]
Table 1 MADS-box genes and their functions
| 基因 Gene | 功能 Function | 表达模式 Expression pattern | 同源基因 Homologous gene |
|---|---|---|---|
| FLC | 控制开花时间,开花抑制子 Control of flowering time, flowering suppressor | 除了茎尖,花前广泛存在于各组织,表达水平下降引起开花 Except shoot apex, widely expressed before flowering, download expression caused flowering | 禾谷类植物(小麦、大麦等)没有FLC类同源MADS-box基因 Cereal plants (wheat, barley, etc) have no FLC homologous genes |
| SVP | 控制开花时间,开花抑制子 Control of flowering time, flowering suppressor | 花前主要存在于花序、花芽、叶的顶端分生组织 Apical meristem of inflorescence, buds, leaves before flowering | 大麦Barley:BM1、BM9、HvVRT2;小麦Wheat:TaVRT2 |
| SOC1 | 控制开花时间 Control of flowering time | 茎尖分生组织、叶、花芽 Shoot apical meristem, leaves, flower buds | 油菜Brassica:LF、MF1、MF2 |
| CAL | 调控花分生组织分化 Floral meristem identity | 花分生组织 Floral meristem | 与AP1是旁系同源 Paralogs of AP1 |
| FUL | 控制开花时间,调控花分生组织分化果实发育 Control of flowering time, floral meristem identity, fruit development | 花序分生组织、胚珠、茎叶 Inflorescence meristem, ovules, cauline leaves | 与AP1是旁系同源 Paralogs of AP1 |
| AGL24 | 控制开花时间,开花激活子 Control of flowering time, flowering activator | 花分生组织 Floral meristem | 与SVP是旁系同源 Paralogs of SVP |
| AP1 | 调控花分生组织分化,A类基因,调控花萼和花瓣形成 Floral meristem identity, A-class homeotic gene, regulated sepals and petals | 整个花分生组织;第1、2轮花器官 Throughout floral meristem, whorls 1 and whorls 2 of floral organs | 金鱼草 Snapdragon:SQUA、DEFH28;水稻Rice:OsMADS14、OsMADS15、OsMADS18 |
| AP3 | B类基因,调控花瓣和雄蕊形成 B-class homeotic gene, regulated petals and stamens | 第2、3轮花器官 Whorls 2 and whorls 3 of floral organs | 金鱼草 Snapdragon:DEF;矮牵牛Petunia:PhGLO1/2;水稻 Rice:OsMADS16;玉米 Maize:Si1 |
| PI | B类基因,调控花瓣和雄蕊形成 B-class homeotic gene, regulated petals and stamens | 第2、3轮花器官 Whorls 2 and whorls 3 of floral organs | 金鱼草Snapdragon:GLO;矮牵牛Petunia:pMADS1、GP;水稻Rice:OsMADS2、OsMADS4 |
| AG | C类基因,调控雄蕊和心皮形成 C-class homeotic gene, regulated stamens and carpels | 第3、4轮花器官 Whorls 3 and whorls 4 of floral organs | 金鱼草Snapdragon:FAR;矮牵牛Petunia:pMADS3;水稻Rice:OsMADS3;玉米Maize:ZAG1 |
| SHP1/2 | D类基因,果实发育和开裂 D-class homeotic gene, fruit development and dehiscence | 胚珠、壳边、果实分裂区域 Ovules, valve margin, fruit dehiscence zone | 金鱼草Snapdragon:PLE;矮牵牛Petunia:FBP6 |
| STK | D类基因,控制胚珠发育 D-class homeotic gene, regulated ovules development | 胚珠 Ovules | 矮牵牛Petunia:FBP7、FBP11;水稻 Rice:OsMADS13、OsMADS21 |
| SEP1/2/3/4 | E类基因,花器官发育辅助因子,调控激活B、C类基因 E-class homeotic gene, co-regulated floral development, activated B-and C-class genes | SEP1/2:4轮花器官;SEP3:第2-3轮花器官;SEP4:第1轮花器官 SEP1/2: all whorls of floral organs; SEP3: whorls 2 and whorls 3; SEP4: whorls 1 | 矮牵牛Petunia:FBP2;番茄Tomato:TM5;水稻Rice:OsMADS1 |
表1 MADS-box基因及其功能[2,14,16-18]
Table 1 MADS-box genes and their functions
| 基因 Gene | 功能 Function | 表达模式 Expression pattern | 同源基因 Homologous gene |
|---|---|---|---|
| FLC | 控制开花时间,开花抑制子 Control of flowering time, flowering suppressor | 除了茎尖,花前广泛存在于各组织,表达水平下降引起开花 Except shoot apex, widely expressed before flowering, download expression caused flowering | 禾谷类植物(小麦、大麦等)没有FLC类同源MADS-box基因 Cereal plants (wheat, barley, etc) have no FLC homologous genes |
| SVP | 控制开花时间,开花抑制子 Control of flowering time, flowering suppressor | 花前主要存在于花序、花芽、叶的顶端分生组织 Apical meristem of inflorescence, buds, leaves before flowering | 大麦Barley:BM1、BM9、HvVRT2;小麦Wheat:TaVRT2 |
| SOC1 | 控制开花时间 Control of flowering time | 茎尖分生组织、叶、花芽 Shoot apical meristem, leaves, flower buds | 油菜Brassica:LF、MF1、MF2 |
| CAL | 调控花分生组织分化 Floral meristem identity | 花分生组织 Floral meristem | 与AP1是旁系同源 Paralogs of AP1 |
| FUL | 控制开花时间,调控花分生组织分化果实发育 Control of flowering time, floral meristem identity, fruit development | 花序分生组织、胚珠、茎叶 Inflorescence meristem, ovules, cauline leaves | 与AP1是旁系同源 Paralogs of AP1 |
| AGL24 | 控制开花时间,开花激活子 Control of flowering time, flowering activator | 花分生组织 Floral meristem | 与SVP是旁系同源 Paralogs of SVP |
| AP1 | 调控花分生组织分化,A类基因,调控花萼和花瓣形成 Floral meristem identity, A-class homeotic gene, regulated sepals and petals | 整个花分生组织;第1、2轮花器官 Throughout floral meristem, whorls 1 and whorls 2 of floral organs | 金鱼草 Snapdragon:SQUA、DEFH28;水稻Rice:OsMADS14、OsMADS15、OsMADS18 |
| AP3 | B类基因,调控花瓣和雄蕊形成 B-class homeotic gene, regulated petals and stamens | 第2、3轮花器官 Whorls 2 and whorls 3 of floral organs | 金鱼草 Snapdragon:DEF;矮牵牛Petunia:PhGLO1/2;水稻 Rice:OsMADS16;玉米 Maize:Si1 |
| PI | B类基因,调控花瓣和雄蕊形成 B-class homeotic gene, regulated petals and stamens | 第2、3轮花器官 Whorls 2 and whorls 3 of floral organs | 金鱼草Snapdragon:GLO;矮牵牛Petunia:pMADS1、GP;水稻Rice:OsMADS2、OsMADS4 |
| AG | C类基因,调控雄蕊和心皮形成 C-class homeotic gene, regulated stamens and carpels | 第3、4轮花器官 Whorls 3 and whorls 4 of floral organs | 金鱼草Snapdragon:FAR;矮牵牛Petunia:pMADS3;水稻Rice:OsMADS3;玉米Maize:ZAG1 |
| SHP1/2 | D类基因,果实发育和开裂 D-class homeotic gene, fruit development and dehiscence | 胚珠、壳边、果实分裂区域 Ovules, valve margin, fruit dehiscence zone | 金鱼草Snapdragon:PLE;矮牵牛Petunia:FBP6 |
| STK | D类基因,控制胚珠发育 D-class homeotic gene, regulated ovules development | 胚珠 Ovules | 矮牵牛Petunia:FBP7、FBP11;水稻 Rice:OsMADS13、OsMADS21 |
| SEP1/2/3/4 | E类基因,花器官发育辅助因子,调控激活B、C类基因 E-class homeotic gene, co-regulated floral development, activated B-and C-class genes | SEP1/2:4轮花器官;SEP3:第2-3轮花器官;SEP4:第1轮花器官 SEP1/2: all whorls of floral organs; SEP3: whorls 2 and whorls 3; SEP4: whorls 1 | 矮牵牛Petunia:FBP2;番茄Tomato:TM5;水稻Rice:OsMADS1 |
图2 花器官发育模型[34,35,36] a,不同层面阐释花发育模型:基因层面为“ABC(D)E模型”——花萼:A类基因调控;花瓣:A+B类基因调控;雄蕊:B+C类基因调控;雌蕊:C类基因单独调控;胚珠:D类基因调控(包括部分C类基因调控);E类基因对所有花器官形成起辅助调控。蛋白层面为“四聚体模型”,由同源或异源二聚体蛋白再聚合形成四聚体——花萼:AP1-SEP-AP1-SEP;花瓣:AP3-PI-SEP-AP1;雄蕊:AP3-PI-SEP-AG;雌蕊:AG-SEP-AG-SEP;胚珠:AG-SEP-SHP-STK。b,单子叶百合科中B类基因覆盖前3轮,导致花萼瓣化。c,毛茛纲部分植物中B类基因只在花瓣中表达。(注: AP2基因不属于MADS-box基因,其他ABCDE类基因均为MADS-box基因。)
Fig.2 Floral organ development models a, Interpreted floral development models at different levels. “The ABC(D)E model” was at genes level. Sepals: A-class genes alone specified; Petals: A-and B-class genes combined to specified; Stamens: B-and C-class genes combined to specified; Carpels: C-class genes alone specified; Ovules: D-class genes specified (partially involved C-class genes); E-class genes were required to co-regulated the formation of floral organs. “The quartet model” was at proteins level. Tetramers were formed by homodimer or heterodimer. Sepals: AP1-SEP-AP1-SEP; Petals: AP3-PI-SEP-AP1; Stamens: AP3-PI-SEP-AG; Carpels: AG-SEP-AG-SEP; Ovules: AG-SEP-SHP-STK. b, Petaloid sepals in Liliaceae of monocotyledon resulted as B-class gene covered the first three whorls. c, B-class genes were expressed only in parts of the petals in some Ranunculus. AP2 gene was not a MADS-box gene, the other ABCDE genes were all MADS-box genes.
图2 花器官发育模型[34,35,36] a,不同层面阐释花发育模型:基因层面为“ABC(D)E模型”——花萼:A类基因调控;花瓣:A+B类基因调控;雄蕊:B+C类基因调控;雌蕊:C类基因单独调控;胚珠:D类基因调控(包括部分C类基因调控);E类基因对所有花器官形成起辅助调控。蛋白层面为“四聚体模型”,由同源或异源二聚体蛋白再聚合形成四聚体——花萼:AP1-SEP-AP1-SEP;花瓣:AP3-PI-SEP-AP1;雄蕊:AP3-PI-SEP-AG;雌蕊:AG-SEP-AG-SEP;胚珠:AG-SEP-SHP-STK。b,单子叶百合科中B类基因覆盖前3轮,导致花萼瓣化。c,毛茛纲部分植物中B类基因只在花瓣中表达。(注: AP2基因不属于MADS-box基因,其他ABCDE类基因均为MADS-box基因。)
Fig.2 Floral organ development models a, Interpreted floral development models at different levels. “The ABC(D)E model” was at genes level. Sepals: A-class genes alone specified; Petals: A-and B-class genes combined to specified; Stamens: B-and C-class genes combined to specified; Carpels: C-class genes alone specified; Ovules: D-class genes specified (partially involved C-class genes); E-class genes were required to co-regulated the formation of floral organs. “The quartet model” was at proteins level. Tetramers were formed by homodimer or heterodimer. Sepals: AP1-SEP-AP1-SEP; Petals: AP3-PI-SEP-AP1; Stamens: AP3-PI-SEP-AG; Carpels: AG-SEP-AG-SEP; Ovules: AG-SEP-SHP-STK. b, Petaloid sepals in Liliaceae of monocotyledon resulted as B-class gene covered the first three whorls. c, B-class genes were expressed only in parts of the petals in some Ranunculus. AP2 gene was not a MADS-box gene, the other ABCDE genes were all MADS-box genes.
图3 MADS-box基因调控花发育网络、花发育过程和相关基因[14,20,21,28,43-45] →表示调控具有促进作用;─┤表示调控具有抑制作用;─表示已经研究确定的调控路径;-----表示未确定的调控路径;*表示该基因不是MADS-box基因;未标记的基本均为MADS-box基因。
Fig.3 MADS-box genes regulate floral development network, floral development processes and related genes →indicated acceleration; ─┤indicated inhibition; ─ represent this regulatory path has been studied;-----represent an unspecified regulatory path; * meant the gene was not a MADS-box gene; Genes unmarked were all MADS-box genes.
图3 MADS-box基因调控花发育网络、花发育过程和相关基因[14,20,21,28,43-45] →表示调控具有促进作用;─┤表示调控具有抑制作用;─表示已经研究确定的调控路径;-----表示未确定的调控路径;*表示该基因不是MADS-box基因;未标记的基本均为MADS-box基因。
Fig.3 MADS-box genes regulate floral development network, floral development processes and related genes →indicated acceleration; ─┤indicated inhibition; ─ represent this regulatory path has been studied;-----represent an unspecified regulatory path; * meant the gene was not a MADS-box gene; Genes unmarked were all MADS-box genes.
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