植物遗传转化中体细胞再生的分子机制及应用研究进展

doi:10.11686/cyxb2023147

草业学报 ›› 2024, Vol. 33 ›› Issue (2): 198-211.DOI: 10.11686/cyxb2023147

• 综合评述 • 上一篇

植物遗传转化中体细胞再生的分子机制及应用研究进展

李玉珠(), 余江弟, 丁菲菲, 苗佳敏, 白小明, 师尚礼()

甘肃农业大学草业学院，草业生态系统教育部重点实验室，中-美草地畜牧业可持续发展研究中心，甘肃兰州 730070

收稿日期:2023-05-08 修回日期:2023-07-07 出版日期:2024-02-20 发布日期:2023-12-12
通讯作者: 师尚礼
作者简介:E-mail： shishl@gsau.edu.cn
李玉珠（1979-），女，陕西宁强人，副教授，博士。E-mail： liyz@gsau.edu.cn
基金资助:
草业生态系统教育部重点实验室（甘肃农业大学）2022年开放课题(KLGE202217);草种创新与草地农业生态系统全国重点实验室开放基金课题(SKLHIGA202301);甘肃农业大学科技创新基金（青年导师扶持基金）(GAU-QDFC-2021-02)

Progress in studies of molecular mechanisms and applications of somatic cell regeneration during genetic transformation

Yu-zhu LI(), Jiang-di YU, Fei-fei DING, Jia-min MIAO, Xiao-ming BAI, Shang-li SHI()

Pratacultural College，Gansu Agricultural University，Key Laboratory of Grassland Ecosystem，Ministry of Education，Sino-U. S. Centers for Grazing Land Ecosystem Sustainability，Lanzhou 730070，China

Received:2023-05-08 Revised:2023-07-07 Online:2024-02-20 Published:2023-12-12
Contact: Shang-li SHI

摘要/Abstract

摘要：

植物遗传转化是转基因技术及以此为基础的基因组编辑、功能基因组学研究和分子育种的关键。其中，物种和基因型差异往往是限制遗传转化效率和基因编辑技术广泛应用的主要瓶颈。随着再生芽发生和体胚发生的分子机制被逐渐探明，在愈伤组织形成、增殖和再生过程中涉及生长素和细胞分裂素合成、响应和信号转导的生长及发育调节基因被用于提高遗传转化效率。本研究首先综述了植物遗传转化过程中体细胞再生的不同途径和方式，以及转化细胞以间接的器官发生方式和体胚发生方式再生的分子机制。然后重点讨论了与生长素和细胞分裂素有关的再生促进基因在提高再生效率，缩短转化时间，以及实现执拗型物种和基因型的遗传转化等方面的应用。最后总结了再生促进基因在转基因和基因编辑中的应用潜力和研究方向。

关键词: 遗传转化, 再生, 再生芽发生, 体胚发生, 再生促进基因

Abstract:

Plant genetic transformation is the key of transgenic technology， genome editing， functional genomics research and molecular breeding. Species and genotype differences are often the main bottlenecks limiting the efficiency of genetic transformation and the wide application of gene editing technology. With the molecular mechanism of de novo shoot organogenesis and somatic embryogenesis being gradually explored， the growth and developmental regulatory genes involved in the synthesis， response and signal transduction of auxin and cytokinin in callus formation， proliferation and regeneration are used to improve genetic transformation efficiency. In this study， we first review the different ways and means of achieving somatic cell regeneration after genetic transformation， and the molecular mechanisms of regeneration for the transformed cells through indirect organogenesis and somatic embryogenesis. Then， the application of regeneration-promoting genes related to auxin and cytokinin in improving regeneration efficiency， shortening transformation time， and realizing genetic transformation of recalcitrant species and genotypes was discussed. Finally， the potential for application of regeneration-promoting genes when working with transgenic and gene-edited cells were summarized and current research directions were discussed.

Key words: genetic transformation, regeneration, do novo shoot organogenesis, somatic embryogenesis, regeneration-promoting genes

李玉珠, 余江弟, 丁菲菲, 苗佳敏, 白小明, 师尚礼. 植物遗传转化中体细胞再生的分子机制及应用研究进展[J]. 草业学报, 2024, 33(2): 198-211.

Yu-zhu LI, Jiang-di YU, Fei-fei DING, Jia-min MIAO, Xiao-ming BAI, Shang-li SHI. Progress in studies of molecular mechanisms and applications of somatic cell regeneration during genetic transformation[J]. Acta Prataculturae Sinica, 2024, 33(2): 198-211.

图/表 1

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基因名称 Gene name	表达盒 Gene cassette	转化物种 Transformed species	外植体 Explant	再生方式 Regeneration	转化效果 Transform efficiency	表型 Phenotype	参考文献 Reference
干细胞关键调节基因Key regulator gene of stem cell
WUSCHEL （WUS）	35S：AtWUS	陆地棉Gossypium hirsutum	下胚轴 Hypocotyl	体胚发生 Embryogenesis	胚性组织显著发生The percentage of explants giving rise to embryogenic tissues was significantly higher than in the control	异常 Abnormal	［54］
	P_G10–90：XVE-O^LexA-min35S：AtWUS	烟草N. tabacum	叶片Leaf	器官发生 Organogenesis	促进转化的顽拗型品种增殖Improving micropropagation in recalcitrant species	异常 Abnormal	［55］
	P_G10–90：XVE-O^LexA-min35S：AtWUS	中粒咖啡Coffea canephora	叶片Leaf	体胚发生 Embryogenesis	提高体胚发生产量400% A 400% increases of somatic embryo production	异常 Abnormal	［56］
	ZmPLTP：ZmWUS2	玉米Z. mays	未成熟胚 Immature embryo	体胚发生 Embryogenesis	促进仅含标记基因的T₀代转基因植物体胚形成和再生Stimulated somatic embryo formation and regeneration of stable T₀ plants that contained the selectable marker	正常 Normal	［59］
生长素相关基因Genes related with auxin
BABY BOOM （BBM）	35S：BnBBM HaUbi：BnBBM	欧洲油菜B. napus	小孢子Microspore	体胚发生Embryogenesis	外植体在无激素的培养基中再生Hormone-free regeneration of explants	异常 Abnormal	［60］
	35S：AtBBM； 35S：BnBBM	烟草N. tabacum	叶片Leaf	器官发生 Organogenesis	转化系自发形成芽和根Transformed lines exhibited spontaneous shoot and root formation	异常 Abnormal	［62］
	35S：TcBBM	可可T. cacao	子叶Cotyledon	体胚发生 Embryogenesis	不需要激素的体胚发生和增殖Increasing SE formation and proliferation without use of exogenous hormones	异常 Abnormal	［63］
	35S：TcBBM-GR	可可T. cacao	子叶Cotyledon	体胚发生 Embryogenesis	赋予叶片体胚发生能力Confering an embryogenic potential to leaves	正常 Normal	［64］
	35S：BnBBM-GR	辣椒C. annuum	子叶Cotyledon	体胚发生 Embryogenesis	执拗型品种高效再生转基因植物Efficiently regenerating transgenic plants from recalcitrant varieties	异常 Abnormal	［65］
LEAFY COTYLEDON1 （LEC1）	35S： PaHAP3A；P_G10–90：XVE-O^LexA-min35S：PaHAP3A	欧洲云杉P. abies	胚性细胞系Embryonic cell lines	体胚发生Embryogenesis	促进体胚分化Leading to the differentiation of ectopic embryos from maturing somatic embryos	正常 Normal	［66］
AGAMOUS-LIKE 15 （AGL15）	35S：gGmAGL15；35S：cGmAGL15	大豆G. max	子叶Cotyledon	体胚发生Embryogenesis	促进体胚发育Enhancing somatic embryo development	异常 Abnormal	［67］
WUS+BBM	Nos：ZmWUS2+ ZmUbi：ZmBBM	玉米Z. mays	未成熟胚 Immature embryo	体胚发生 Embryogenesis	实现多个执拗型品种高频转化High transformation frequencies in numerous previously nontransformable maize inbred lines	正常 Normal	［69］
	Nos：ZmWUS2+ ZmUbi：ZmBBM	高粱S. bicolor 甘蔗S. officinarum 水稻O. sativa	未成熟胚 Immature embryo	体胚发生 Embryogenesis	促进高粱未成熟胚、甘蔗愈伤组织和籼稻愈伤组织的转化Stimulating transformation in sorghum immature embryos， sugarcane callus， and indica rice callus	正常 Normal	［69］
	Nos：ZmWUS2+ ZmUbi：ZmBBM	玉米Z. mays 高粱S. bicolor	未成熟胚 Immature embryo	体胚发生 Embryogenesis	不需要选择性标记基因获得执拗型玉米自交系B73和高粱P898012的有效转化Efficient transformation of recalcitrant maize inbred B73 and sorghum P898012 genotypes without use of a selectable marker gene	正常 Normal	［70］
WUS+BBM	Nos：ZmWUS2+ZmPLTP：ZmBBM	玉米Z. mays	未成熟胚 Immature embryo	体胚发生 Embryogenesis	绕过愈伤组织诱导直接产生体胚并再生Somatic embryos rapidly formed and directly germinated into plants without a callus phase	正常 Normal	［72］
	ZmAxig1：ZmWUS2+ZmPLTP：ZmBBM	玉米Z. mays	未成熟胚 Immature embryo	体胚发生 Embryogenesis	绕过愈伤组织诱导直接产生体胚并再生，且T₁代种子正常萌发Somatic embryos rapidly formed and directly germinated into plants without a callus phase， and single-copy T₁ seed germinated normally	正常 Normal	［72］
	ZmPLTP：ZmWUS2+ZmPLTP：ZmBBM	高粱S. bicolor	未成熟胚 Immature embryo	体胚发生 Embryogenesis	更多基因型被转化，并缩短转基因植株再生时间More sorghum genotypes amenable to transformation and a faster time to generate transformed plants	正常 Normal	［73］
	Nos：ZmWUS2-3xEnhZmUbi：ZmBBM	玉米Z. mays 高粱S. bicolor	叶片Leaf	体胚发生 Embryogenesis	通过体胚直接发生而显著提高玉米和高粱的转化效率并获得Cas9介导的基因编辑植株Substantially improving leaf transformation in maize and sorghum by direct SE， allowing the recovery of plants with Cas9-mediated gene dropouts and targeted gene insertion	正常 Normal	［74］
	Nos：ZmWUS2-3xEnhZmUbi：ZmBBM	苔麸Eragrostis tef；柳枝稷Panicum virgatum；珍珠粟Pennisetum glaucum；梁Setaria italica；黑麦Secale cereale；大麦Hordeum vulgare；水稻O. sativa	叶片Leaf	体胚发生 Embryogenesis	获得禾本科4个不同亚科7个物种的体胚再生转化植株Embryogenic callus and regenerated plantlets were successfully produced in seven species spanning four grass subfamilies	正常 Normal	［74］
CK相关基因CK related genes
CUP-SHAPED COTYLEDON1 or 2CUC1 or 2	35S：AtCUC1 or 2	拟南芥A. thaliana	幼苗Seedling	器官发生 Organogenesis	转化愈伤组织不定芽数量提高10倍The average number of adventitious shoots on a callus was 10 times than that in the control	异常 Abnormal	［31］
ENHANCER OF SHOOT REGENERATION2 （ESR2）	P_G10–90：XVE-O^LexA-min35S：AtESR2	拟南芥A. thaliana	根Root	器官发生 Organogenesis	不需要添加外源CK而促进芽再生Promoted shoot regeneration independently of added cytokinin	正常 Normal	［34］
Knotted1 （Kn1）	35S：ZmKn1	甜橙Citrus sinensis	茎段Internodal stem segments	器官发生 Organogenesis	转化效率提高3~15倍Enhances transformation efficiencies from 3 to 15 fold	异常 Abnormal	［75］
Knotted1 （Kn1）	35S：ZmKn1	烟草N. tabacum	叶片Leaf	器官发生Organogenesis	转化效率提高3倍3-fold increase in transformation efficiency	异常Abnormal	［76］
MONOPTEROS （MP）	AtMP：AtMPΔ	拟南芥A. thaliana	根Root 叶Leaf 叶柄Petiole 子叶Cotyledon	器官发生 Organogenesis	促进再生芽发生Promote de novo shoot organogenesis	异常Abnormal	［77］
GROWTH-REGULATING FACTOR5（GRF5）	2×35S：AtGRF5；2×35S：BvGRF5-L	甜菜Beta vulgaris	下胚轴Cotyledon 子叶Hypocotyl	器官发生 Organogenesis	提高转化效率和再生效率，包括执拗型品种Accelerated shoot organogenesis and resulted in significant improvements in transformation， including recalcitrant varieties	正常 Normal	［78］
	2×35S：AtGRF5；2×35 S：HaGRF5-L	向日葵Helianthus annuu	子叶Cotyledon	器官发生Organogenesis	促进转化细胞中芽的产生Increasing in the production of developing transgenic shoots	正常Normal	［78］
	PcUbi4-2：： GmGRF5-L	大豆G. max	茎节Primary node	器官发生Organogenesis	促进转化细胞中芽的产生Increasing in the production of developing transgenic shoots	正常 Normal	［78］
	PcUbi4-2： BnGRF5-L	欧洲油菜B. napus	下胚轴Hypocotyl	器官发生Organogenesis	显著提高转化效率Significantly increases in genetic transformation of the explant tissue	正常 Normal	［78］
	BdEF1：AtGRF5；BdEF 1：ZmGRF5-L1/2	玉米Z. mays	未成熟胚Immature embryo	体胚发生Embryogenesis	促进胚性愈伤组织增殖，提高转化效率A higher rate of embryogenic callus growth leading to an increased recovery of transgenic plants	正常 Normal	［78］
GRF-INTERACTING FACTOR （GRF-FIF）	TaUbi：GRF4-GIF1	小麦T. aestivum	未成熟胚Immature embryo	器官发生Organogenesis	平均再生效率比对照高7.8倍The average regeneration efficiency of the GRF4-GIF1 chimera was 7.8-fold higher than the empty vector control	正常 Normal	［79］
	TaUbi：GRF4-GIF1	水稻O. sativa	种子Seed	器官发生Organogenesis	转化愈伤组织的再生效率比对照提高2.1倍A 2.1-fold increase in regeneration efficiency in the calli transformed compared with those transformed with the control vectors	正常 Normal	［79］
	ClUbi：GRF4-GIF1	柠檬Citrus×limon	上胚轴Etiolated epicotyl	器官发生 Organogenesis	再生效率比对照提高4.7倍A 4.7-fold increase in regeneration frequency relative to those transformed with the empty vector control	正常 Normal	［79］
ISOPENTENYL TRANSFERASE （IPT）	35S：ipt	烟草N. tabacum	茎Stem 叶片Leaf	器官发生Organogenesis	芽数量增加2.4倍，再生时间缩短1.5~2.3倍2.4-fold increase in the number of shoots formation and 1.5-2.3 fold decrease in the time of shoots formation	异常 Abnormal	［80-81］
	35S：ipt	黄瓜Cucumis sativus	幼苗Seedling	器官发生Organogenesis	芽平均大小增加7.5倍7.5-fold increase in the size of shoots	异常 Abnormal	［80-81］
	35S：ipt in Ac	烟草N. tabacum	叶片Leaf	器官发生Organogenesis	转基因植株的选择标记基因被自动切除Marker-free phenotypically normal transgenic plants	正常 Normal	［82］
	35S：ipt in Ac	杂交山杨Populus sieboldii×Populus grandidentata	茎Stem	器官发生Organogenesis	转基因植株的选择标记基因被自动切除Marker-free phenotypically normal transgenic plants	正常 Normal	［82］
	35S：GVG- 6×UAS-min3-5S：ipt	烟草N. tabacum	叶片Leaf	器官发生Organogenesis	Dex诱导系统控制下，再生植株增加24.3倍A 24.3-fold increase in shoots regenerants under the control of the Dex-inducible system	正常 Normal	［83］
	35S：GVG - 6×UAS-min3-5S：ipt	莴苣Lactuca sativa	子叶Cotyledon	器官发生Organogenesis	Dex诱导系统控制下，再生植株增加6.6倍A 6.6-fold increase in shoots regenerants under the control of the Dex-inducible system	正常 Normal	［83］
	Nos：ZmWUS 2-ZmUbi：IPT or Nos：ZmW US2- ZmUbi：AtSTM	拟南芥A. thaliana 烟草N. tabacum	幼苗Seedling	器官发生Organogenesis	无须组织培养，快速产生基因编辑植株Generating gene-edited plants through de novo meristem induction	正常 Normal	［84］
	Nos：ZmWUS 2-ZmUbi：IPT or Nos：ZmW US2- ZmUbi：AtSTM	番茄Solanum lycopersicum；马铃薯S. tuberosum；葡萄Vitis vinifera	幼苗Seedling	器官发生Organogenesis	无须组织培养，快速产生基因编辑植株Generating gene-edited plants through de novo meristem induction	正常 Normal	［84］

基因名称 Gene name	表达盒 Gene cassette	转化物种 Transformed species	外植体 Explant	再生方式 Regeneration	转化效果 Transform efficiency	表型 Phenotype	参考文献 Reference
干细胞关键调节基因Key regulator gene of stem cell
WUSCHEL （WUS）	35S：AtWUS	陆地棉Gossypium hirsutum	下胚轴 Hypocotyl	体胚发生 Embryogenesis	胚性组织显著发生The percentage of explants giving rise to embryogenic tissues was significantly higher than in the control	异常 Abnormal	［54］
	P_G10–90：XVE-O^LexA-min35S：AtWUS	烟草N. tabacum	叶片Leaf	器官发生 Organogenesis	促进转化的顽拗型品种增殖Improving micropropagation in recalcitrant species	异常 Abnormal	［55］
	P_G10–90：XVE-O^LexA-min35S：AtWUS	中粒咖啡Coffea canephora	叶片Leaf	体胚发生 Embryogenesis	提高体胚发生产量400% A 400% increases of somatic embryo production	异常 Abnormal	［56］
	ZmPLTP：ZmWUS2	玉米Z. mays	未成熟胚 Immature embryo	体胚发生 Embryogenesis	促进仅含标记基因的T₀代转基因植物体胚形成和再生Stimulated somatic embryo formation and regeneration of stable T₀ plants that contained the selectable marker	正常 Normal	［59］
生长素相关基因Genes related with auxin
BABY BOOM （BBM）	35S：BnBBM HaUbi：BnBBM	欧洲油菜B. napus	小孢子Microspore	体胚发生Embryogenesis	外植体在无激素的培养基中再生Hormone-free regeneration of explants	异常 Abnormal	［60］
	35S：AtBBM； 35S：BnBBM	烟草N. tabacum	叶片Leaf	器官发生 Organogenesis	转化系自发形成芽和根Transformed lines exhibited spontaneous shoot and root formation	异常 Abnormal	［62］
	35S：TcBBM	可可T. cacao	子叶Cotyledon	体胚发生 Embryogenesis	不需要激素的体胚发生和增殖Increasing SE formation and proliferation without use of exogenous hormones	异常 Abnormal	［63］
	35S：TcBBM-GR	可可T. cacao	子叶Cotyledon	体胚发生 Embryogenesis	赋予叶片体胚发生能力Confering an embryogenic potential to leaves	正常 Normal	［64］
	35S：BnBBM-GR	辣椒C. annuum	子叶Cotyledon	体胚发生 Embryogenesis	执拗型品种高效再生转基因植物Efficiently regenerating transgenic plants from recalcitrant varieties	异常 Abnormal	［65］
LEAFY COTYLEDON1 （LEC1）	35S： PaHAP3A；P_G10–90：XVE-O^LexA-min35S：PaHAP3A	欧洲云杉P. abies	胚性细胞系Embryonic cell lines	体胚发生Embryogenesis	促进体胚分化Leading to the differentiation of ectopic embryos from maturing somatic embryos	正常 Normal	［66］
AGAMOUS-LIKE 15 （AGL15）	35S：gGmAGL15；35S：cGmAGL15	大豆G. max	子叶Cotyledon	体胚发生Embryogenesis	促进体胚发育Enhancing somatic embryo development	异常 Abnormal	［67］
WUS+BBM	Nos：ZmWUS2+ ZmUbi：ZmBBM	玉米Z. mays	未成熟胚 Immature embryo	体胚发生 Embryogenesis	实现多个执拗型品种高频转化High transformation frequencies in numerous previously nontransformable maize inbred lines	正常 Normal	［69］
	Nos：ZmWUS2+ ZmUbi：ZmBBM	高粱S. bicolor 甘蔗S. officinarum 水稻O. sativa	未成熟胚 Immature embryo	体胚发生 Embryogenesis	促进高粱未成熟胚、甘蔗愈伤组织和籼稻愈伤组织的转化Stimulating transformation in sorghum immature embryos， sugarcane callus， and indica rice callus	正常 Normal	［69］
	Nos：ZmWUS2+ ZmUbi：ZmBBM	玉米Z. mays 高粱S. bicolor	未成熟胚 Immature embryo	体胚发生 Embryogenesis	不需要选择性标记基因获得执拗型玉米自交系B73和高粱P898012的有效转化Efficient transformation of recalcitrant maize inbred B73 and sorghum P898012 genotypes without use of a selectable marker gene	正常 Normal	［70］
WUS+BBM	Nos：ZmWUS2+ZmPLTP：ZmBBM	玉米Z. mays	未成熟胚 Immature embryo	体胚发生 Embryogenesis	绕过愈伤组织诱导直接产生体胚并再生Somatic embryos rapidly formed and directly germinated into plants without a callus phase	正常 Normal	［72］
	ZmAxig1：ZmWUS2+ZmPLTP：ZmBBM	玉米Z. mays	未成熟胚 Immature embryo	体胚发生 Embryogenesis	绕过愈伤组织诱导直接产生体胚并再生，且T₁代种子正常萌发Somatic embryos rapidly formed and directly germinated into plants without a callus phase， and single-copy T₁ seed germinated normally	正常 Normal	［72］
	ZmPLTP：ZmWUS2+ZmPLTP：ZmBBM	高粱S. bicolor	未成熟胚 Immature embryo	体胚发生 Embryogenesis	更多基因型被转化，并缩短转基因植株再生时间More sorghum genotypes amenable to transformation and a faster time to generate transformed plants	正常 Normal	［73］
	Nos：ZmWUS2-3xEnhZmUbi：ZmBBM	玉米Z. mays 高粱S. bicolor	叶片Leaf	体胚发生 Embryogenesis	通过体胚直接发生而显著提高玉米和高粱的转化效率并获得Cas9介导的基因编辑植株Substantially improving leaf transformation in maize and sorghum by direct SE， allowing the recovery of plants with Cas9-mediated gene dropouts and targeted gene insertion	正常 Normal	［74］
	Nos：ZmWUS2-3xEnhZmUbi：ZmBBM	苔麸Eragrostis tef；柳枝稷Panicum virgatum；珍珠粟Pennisetum glaucum；梁Setaria italica；黑麦Secale cereale；大麦Hordeum vulgare；水稻O. sativa	叶片Leaf	体胚发生 Embryogenesis	获得禾本科4个不同亚科7个物种的体胚再生转化植株Embryogenic callus and regenerated plantlets were successfully produced in seven species spanning four grass subfamilies	正常 Normal	［74］
CK相关基因CK related genes
CUP-SHAPED COTYLEDON1 or 2CUC1 or 2	35S：AtCUC1 or 2	拟南芥A. thaliana	幼苗Seedling	器官发生 Organogenesis	转化愈伤组织不定芽数量提高10倍The average number of adventitious shoots on a callus was 10 times than that in the control	异常 Abnormal	［31］
ENHANCER OF SHOOT REGENERATION2 （ESR2）	P_G10–90：XVE-O^LexA-min35S：AtESR2	拟南芥A. thaliana	根Root	器官发生 Organogenesis	不需要添加外源CK而促进芽再生Promoted shoot regeneration independently of added cytokinin	正常 Normal	［34］
Knotted1 （Kn1）	35S：ZmKn1	甜橙Citrus sinensis	茎段Internodal stem segments	器官发生 Organogenesis	转化效率提高3~15倍Enhances transformation efficiencies from 3 to 15 fold	异常 Abnormal	［75］
Knotted1 （Kn1）	35S：ZmKn1	烟草N. tabacum	叶片Leaf	器官发生Organogenesis	转化效率提高3倍3-fold increase in transformation efficiency	异常Abnormal	［76］
MONOPTEROS （MP）	AtMP：AtMPΔ	拟南芥A. thaliana	根Root 叶Leaf 叶柄Petiole 子叶Cotyledon	器官发生 Organogenesis	促进再生芽发生Promote de novo shoot organogenesis	异常Abnormal	［77］
GROWTH-REGULATING FACTOR5（GRF5）	2×35S：AtGRF5；2×35S：BvGRF5-L	甜菜Beta vulgaris	下胚轴Cotyledon 子叶Hypocotyl	器官发生 Organogenesis	提高转化效率和再生效率，包括执拗型品种Accelerated shoot organogenesis and resulted in significant improvements in transformation， including recalcitrant varieties	正常 Normal	［78］
	2×35S：AtGRF5；2×35 S：HaGRF5-L	向日葵Helianthus annuu	子叶Cotyledon	器官发生Organogenesis	促进转化细胞中芽的产生Increasing in the production of developing transgenic shoots	正常Normal	［78］
	PcUbi4-2：： GmGRF5-L	大豆G. max	茎节Primary node	器官发生Organogenesis	促进转化细胞中芽的产生Increasing in the production of developing transgenic shoots	正常 Normal	［78］
	PcUbi4-2： BnGRF5-L	欧洲油菜B. napus	下胚轴Hypocotyl	器官发生Organogenesis	显著提高转化效率Significantly increases in genetic transformation of the explant tissue	正常 Normal	［78］
	BdEF1：AtGRF5；BdEF 1：ZmGRF5-L1/2	玉米Z. mays	未成熟胚Immature embryo	体胚发生Embryogenesis	促进胚性愈伤组织增殖，提高转化效率A higher rate of embryogenic callus growth leading to an increased recovery of transgenic plants	正常 Normal	［78］
GRF-INTERACTING FACTOR （GRF-FIF）	TaUbi：GRF4-GIF1	小麦T. aestivum	未成熟胚Immature embryo	器官发生Organogenesis	平均再生效率比对照高7.8倍The average regeneration efficiency of the GRF4-GIF1 chimera was 7.8-fold higher than the empty vector control	正常 Normal	［79］
	TaUbi：GRF4-GIF1	水稻O. sativa	种子Seed	器官发生Organogenesis	转化愈伤组织的再生效率比对照提高2.1倍A 2.1-fold increase in regeneration efficiency in the calli transformed compared with those transformed with the control vectors	正常 Normal	［79］
	ClUbi：GRF4-GIF1	柠檬Citrus×limon	上胚轴Etiolated epicotyl	器官发生 Organogenesis	再生效率比对照提高4.7倍A 4.7-fold increase in regeneration frequency relative to those transformed with the empty vector control	正常 Normal	［79］
ISOPENTENYL TRANSFERASE （IPT）	35S：ipt	烟草N. tabacum	茎Stem 叶片Leaf	器官发生Organogenesis	芽数量增加2.4倍，再生时间缩短1.5~2.3倍2.4-fold increase in the number of shoots formation and 1.5-2.3 fold decrease in the time of shoots formation	异常 Abnormal	［80-81］
	35S：ipt	黄瓜Cucumis sativus	幼苗Seedling	器官发生Organogenesis	芽平均大小增加7.5倍7.5-fold increase in the size of shoots	异常 Abnormal	［80-81］
	35S：ipt in Ac	烟草N. tabacum	叶片Leaf	器官发生Organogenesis	转基因植株的选择标记基因被自动切除Marker-free phenotypically normal transgenic plants	正常 Normal	［82］
	35S：ipt in Ac	杂交山杨Populus sieboldii×Populus grandidentata	茎Stem	器官发生Organogenesis	转基因植株的选择标记基因被自动切除Marker-free phenotypically normal transgenic plants	正常 Normal	［82］
	35S：GVG- 6×UAS-min3-5S：ipt	烟草N. tabacum	叶片Leaf	器官发生Organogenesis	Dex诱导系统控制下，再生植株增加24.3倍A 24.3-fold increase in shoots regenerants under the control of the Dex-inducible system	正常 Normal	［83］
	35S：GVG - 6×UAS-min3-5S：ipt	莴苣Lactuca sativa	子叶Cotyledon	器官发生Organogenesis	Dex诱导系统控制下，再生植株增加6.6倍A 6.6-fold increase in shoots regenerants under the control of the Dex-inducible system	正常 Normal	［83］
	Nos：ZmWUS 2-ZmUbi：IPT or Nos：ZmW US2- ZmUbi：AtSTM	拟南芥A. thaliana 烟草N. tabacum	幼苗Seedling	器官发生Organogenesis	无须组织培养，快速产生基因编辑植株Generating gene-edited plants through de novo meristem induction	正常 Normal	［84］
	Nos：ZmWUS 2-ZmUbi：IPT or Nos：ZmW US2- ZmUbi：AtSTM	番茄Solanum lycopersicum；马铃薯S. tuberosum；葡萄Vitis vinifera	幼苗Seedling	器官发生Organogenesis	无须组织培养，快速产生基因编辑植株Generating gene-edited plants through de novo meristem induction	正常 Normal	［84］

[1]	姜凯, 吴雪莉, 刘奕君, 马越, 宋洋, 卢文杰, 王增裕. 海滨雀稗以hpt与bar基因为筛选标记的转化体系比较研究[J]. 草业学报, 2023, 32(1): 165-177.
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植物遗传转化中体细胞再生的分子机制及应用研究进展

Progress in studies of molecular mechanisms and applications of somatic cell regeneration during genetic transformation

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