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中国生物工程杂志

CHINA BIOTECHNOLOGY
中国生物工程杂志
中国生物工程杂志  2022, Vol. 42 Issue (8): 85-98    DOI: 10.13523/j.cb.2204042
综述     
促进植物高效遗传转化的发育调节因子及其在玉米中的应用进展*
何伟1,2(),朱美睿1,2,单海艳1,2,江易林1,2,安学丽1,2,3(),万向元1,2,3()
1.北京科技大学生物与农业研究中心 化学与生物工程学院 顺德研究生院 北京 100083
2.北京中智生物农业国际研究院 北京 100192
3.北京首佳利华科技有限公司 主要作物生物育种北京市工程实验室 生物育种北京市国际科技合作基地 北京 100192
Development Regulatory Factors Promoting Efficient Plant Genetic Transformation and Their Application in Maize
HE Wei1,2(),ZHU Mei-rui1,2,SHAN Hai-yan1,2,JIANG Yi-lin1,2,AN Xue-li1,2,3(),WAN Xiang-yuan1,2,3()
1. Research Center of Biology and Agriculture, Shunde Graduate School, School of Chemistry and Biological Engineering, University of Science and Technology Beijing, Beijing 100083, China
2. Zhongzhi International Institute of Agricultural Biosciences, Beijing 100192, China
3. Beijing Engineering Laboratory of Main Crop Bio-Tech Breeding, Beijing International Science and Technology Cooperation Base of Bio-Tech Breeding, Beijing Solidwill Sci-Tech Co., Ltd., Beijing 100192, China
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摘要：

高效遗传转化技术体系的建立对植物功能基因组学研究和作物新品种的培育均具有促进作用,目前,再生效率低下是限制许多植物高效遗传转化体系建立的主要技术屏障之一。随着对植物分生组织和体细胞胚形成过程研究的深入,鉴定到了一些关键调控基因,统称为发育调节因子。发育调节因子应用于植物遗传转化后,可以有效改善植物分生组织诱导和再生能力,为提高遗传转化效率提供了重要机遇。综述了7类发育调节因子在提高植物遗传转化效率中的研究进展,重点介绍了其中3类在促进玉米遗传转化中的应用,最后展望了建立植物高效遗传转化体系的发展方向。
关键词： 植物;  玉米;  发育调节因子;  遗传转化;  再生能力    
Abstract:

The establishment of efficient genetic transformation systems can promote the study of plant functional genomics and the cultivation of new crop varieties. At present, low regeneration efficiency is one of the main technical barriers to the establishment of efficient genetic transformation systems in many plants. With the deepening of research on plant meristem and somatic embryo formation, some key regulatory genes have been identified, which are collectively called developmental regulators. The application of developmental regulatory factors in plant genetic transformation can effectively improve the establishment of plant meristem induction and regeneration ability, which provides an important opportunity to improve the efficiency of genetic transformation. In this paper, the research progress of seven types of developmental regulatory factors in improving the efficiency of plant genetic transformation was reviewed, with emphasis on the application of three types of developmental regulatory factors in promoting maize genetic transformation. Finally, the development direction of establishing efficient plant genetic transformation system was presented.
Key words: Plant    Maize    Developmental regulators    Genetic transformation    Regenerative capacity
收稿日期: 2022-04-18 出版日期: 2022-09-07
ZTFLH:  Q819  
基金资助: * 国家自然科学基金面上项目(31971958);中央高校基本科研业务费专项资金(06500060)
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何伟
朱美睿
单海艳
江易林
安学丽
万向元

引用本文:

何伟,朱美睿,单海艳,江易林,安学丽,万向元. 促进植物高效遗传转化的发育调节因子及其在玉米中的应用进展*[J]. 中国生物工程杂志, 2022, 42(8): 85-98.

HE Wei,ZHU Mei-rui,SHAN Hai-yan,JIANG Yi-lin,AN Xue-li,WAN Xiang-yuan. Development Regulatory Factors Promoting Efficient Plant Genetic Transformation and Their Application in Maize. China Biotechnology, 2022, 42(8): 85-98.

链接本文:

https://manu60.magtech.com.cn/biotech/CN/10.13523/j.cb.2204042    或    https://manu60.magtech.com.cn/biotech/CN/Y2022/V42/I8/85

图1  发育调节因子SERKs、LEC和WIND1促进植株转化和再生功能示意图
种类 发育调节因子 功能 基因名称 应用植物 转化提升效果 参考文献
1 SOMATIC EMBRYOGENESIS RECEPTOR-LIKE KINASE (SERK) 编码的体细胞胚发生受体类激酶通过信号级联放大促进体细胞胚发生早期基因的表达 AtSERK1 拟南芥 使体细胞胚发生的能力增加3~4 倍 [11]
CcSERK1 咖啡 使体细胞胚数量增加2倍 [12]
2 LEAFY COTYLEDON1(LEC1) 编码的转录因子同源物(CCAAT盒结合因子HAP3亚基)诱导营养细胞中胚胎特异性基因的表达并促进体细胞胚样结构的形成 AtLEC1 拟南芥 — [13]
LEAFY COTYLEDON2 (LEC2) 编码的B3结构域转录因子在胚胎发生中具有重要作用 AtLEC2 拟南芥 — [14]
TcLEC2 可可树 体细胞胚生成效率提升2倍 [15]
3 WIND1 编码的AP2/ERF 转录因子通过ARR依赖性信号通路促进细胞去分化从而获得细胞增殖能力 AtWIND1 拟南芥、
油菜、番
茄和烟草		 可绕过伤口和生长素预处理促进愈伤组织形成 [16⇓-18]
4 BABY BOOM (BBM) 和WUSCHEL (WUS) BABY BOOM(BBM)编码的APETALA2 (AP2) 家族/乙烯反应元件结合因子 (AP2/ERF) 结构域转录因子通过促进细胞分裂和去分化的信号通路来增强体细胞胚发生;
WUSCHEL(WUS)编码的同源盒转录因子通过WUS-CLV反馈系统维持干细胞库的稳定和顶端分生组织的发育		 ZmBbm;OsBbm;
ZmWus2		 玉米、高粱、
甘蔗和水稻		 实现顽固品种的转化和/或提高转化效率 [19]
AtWUS 苜蓿 可在无植物激素的再生系统中提高再生和转化效率 [20]
HvBBM;HvWUS 大麦 转化效率提升3倍多 [21]
BnBBM 辣椒 转化效率达到1%左右 [22-23]
AtBBM 烟草 增强了烟草的再生能力 [24]
TcBBM 可可树 显著增强了体细胞胚的增殖 [25]
AtWUS 棉花 显著提高胚性愈伤组织形成效率 [26-27]
AtWUS 咖啡 体细胞胚生成产量提升400% [28]
5 GROWTH-REGULATING FACTOR (GRF) 和 GRF-INTERACTING FACTOR (GIF) 生长调节因子(GRF)与转录辅因子GRF-INTERACTING FACTOR(GIF)相互作用并形成功能性转录复合体,然后调节干细胞与其快速分裂的子细胞之间的过渡,从而促进细胞增殖 TaGRF4;TaGIF1 小麦、水稻 再生效率分别提高7.8倍和2.1倍 [29]
ClGRF4;ClGIF1;
VvGRF4;VvGIF1		 柑橘 再生效率提高4.7 倍
ZmGRF5-LIKE1;
ZmGRF5-LIKE2		 玉米 提高转化效率约3倍 [30]
AtGRF5;
BvGRF5-LIKE		 甜菜 实现了顽固品种的转化
GmGRF5-LIKE 大豆 促进转基因芽形成
AtGRF5;
HaGRF5-LIKE		 向日葵 促进转基因芽形成
BnGRF5-LIKE 油菜 促进愈伤组织形成
ClGRF4;ClGIF1 西瓜 提高转化效率约9倍 [31]
6 WOX5 WUSCHEL家族基因 TaWOX5 小黑麦、黑麦、
大麦和玉米		 大幅度提高了小麦等植物的转化效率 [32]
表1  促进单双子叶植物高效遗传转化的发育调节因子
图2  发育调节因子BBM和WUS促进细胞去分化和体细胞胚形成功能示意图
图3  GRF-GIF复合体、WOX基因和LBD基因促进植株转化功能示意图
基因名称 玉米自交系 农杆菌 形态发生因子
表达盒		 形态发生因子
切除方式		 优点 缺点 参考文献
ZmBBm,
ZmWus2		 PHH5G
PHP38
PHN46
PH581
PH0AZ		 LBA4404
(THY-)		 Ubipro:BBm:pinII
Nospro:Wus2:pinII		 干燥诱导启动子
Rab17pro控制CRE/loxP的切除系统		 显著提高了多个玉米自交系的遗传转化效率,包括顽固基因型自交系 形态发生因子组成型表达,当切除不彻底时,会导致无法获得健康的T0植株 [19]
ZmBBm,
ZmWus2		 B73 AGL1 Ubipro:BBm:nos
Nospro:Wus2: nos		 干燥诱导启动子
Rab17pro控制CRE/loxP的切除系统		 提高了顽固基因型玉米自交系B73的转化率,降低了基因型依赖性 同上 [47]
ZmBBm,
ZmWus2		 B73
Mo17
PHH5G
PH1V69
PHR03
PH184C
PH1V5T		 LBA4404
(THY-)		 Zm-PLTPpro:BBm:
Os-T28
ZmAxig1pro:Wus2:
IN2-1		 无切除 新的启动子使BBM不在根中表达,提高了T0健康植株的数目;绕过愈伤组织可直接形成健康的T0植株 过表达两种形态发生因子会导致副效应过大,同时消除难度较大,对T0植株的健康有不利影响 [86]
ZmBBm,
ZmWus2		 ND88 EHA105 Ubipro:BBm:
Wus2:HSPt		 干燥诱导启动子
Rab17pro控制CRE/loxP的切除系统		 构建一个三元载体系统,结合WUS和BBM,进一步增强了转化效率 形态发生因子组成型表达,当切除不彻底时,会导致无法获得健康的T0植株 [50]
ZmWus2 HC69
PH1V69
PHW52
PH4BAH
PH2KD1
PH28SV
PH2Y8G
PH4B9Z		 LBA4404
(THY-)		 Zm-PLTPpro:Wus2:
IN2-1		 无切除 只使用WUS2基因,降低了在转化后、再生前切除形态发生基因的难度;同时添加CRC表达盒使得无须切除系统 无 [49]
表2  形态发生因子ZmBBM和ZmWUS2在玉米遗传转化体系改良中的应用
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