十字花科蔬菜硫代葡萄糖苷的代谢调控、遗传定位及基因工程改良研究进展

doi:10.16420/j.issn.0513-353x.2025-0646

摘要/Abstract

摘要：

硫代葡萄糖苷（glucosinolates，GSL）简称硫苷，是十字花科蔬菜的一类重要次生代谢产物，在十字花科蔬菜的营养品质、抗逆性和加工利用价值中发挥核心作用。本研究中系统综述了十字花科蔬菜硫苷的代谢调控、遗传定位及基因工程改良研究进展，涵盖（1）硫苷的生物学功能、种类与合成途径；（2）硫苷合成、转运、水解的关键调控因子及其调控机制；（3）硫苷的遗传定位研究与基因工程改良的重要成果，以期为十字花科蔬菜硫苷性状的遗传改良提供参考。

关键词: 十字花科, 硫代葡萄糖苷, 遗传定位, 转录因子, QTL定位, 基因工程

Abstract:

Glucosinolates（GSL），commonly referred to as glucosinolates，are an important class of secondary metabolites in cruciferous vegetables and play a central role in their nutritional quality，stress resistance，and processing value. This study systematically reviews the research progress on the metabolic regulation，genetic mapping，and genetic engineering improvement of glucosinolates in cruciferous vegetables，covering（1）the biological functions，types，and biosynthetic pathways of glucosinolates；（2）key regulatory factors and mechanisms involved in their biosynthesis，transport，and hydrolysis；（3）major achievements in genetic mapping studies and genetic engineering improvement of glucosinolates，aiming to provide a reference for the genetic improvement of glucosinolate traits in cruciferous vegetables.

Key words: Brassicaceae, glucosinolate, genetic mapping, transcription factor, QTL mapping, genetic engineering

刘嘉超, 郭尊奕, 杨桂玲, 黎仕骏, 陈琳, 陈长明. 十字花科蔬菜硫代葡萄糖苷的代谢调控、遗传定位及基因工程改良研究进展[J]. 园艺学报, 2026, 53(8): 2502-2528.

LIU Jiachao, GUO Zunyi, YANG Guiling, LI Shijun, CHEN Lin, CHEN Changming. Research Progress on Metabolic Regulation，Genetic Mapping and Genetic Engineering Improvement of Glucosinolates in Cruciferous Vegetables[J]. Acta Horticulturae Sinica, 2026, 53(8): 2502-2528.

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物种 Species	杂交组合 Hybridized combination	群体类型 Group type	关键QTL定位 Key QTL localization	文献 Reference
甘蓝型油菜 Brassica napus L.	GH06 × P174	RIL	qGSL-A03-1、qGSL-A09-2、qGSL-A09-3、qGSL-C02-1、qGSL-A01-11、qGSL-A07-12、qGSL-A09-11、qGSL-C02-12、qGSL-A06-11	荐红举等,2014
	GH06 × 油研2号 GH06 × Youyan 2	RIL	qPRO-10-1、qPRO-14-2、qPRO-24-3、qNAP-3-1、qNAP-10-2、qNAP-24-3、qGBN-9-1、qGBN-20-2、qNAS-9-1、qNAS-15-2、qNAS-21-3	刘水燕等,2014
	EXPRESS × SWU07	DH	qGSLEA04-1、qGSLEA06-1、qGSLEA09-1、qGSLEA09-2	He et al.,2018
	G120 × 9172	DH	qGSL-C2、qGSL-A3	Liu et al.,2020b
	ZS11 × G120	DH	cqSGC-A1-1、cqSGC-A1-2、cqSGC-A1-3、cqSGC-A1-4、cqSGC-A2-1、cqSGC-A2-2、cqSGC-A3、cqSGC-A4、cqSGC-A5、cqSGC-A6-1、cqSGC-A6-2、cqSGC-A7、cqSGC-C2	周显明,2021
	M305 × Z114	DH	qGLS-A02、qGLS-A04、qSCC/GLS-C02r、 qSCC/ADF/NDF/GLS-A04、qSCC/NDF/GLS-C06、qSCC/NDF/GLS-C08、qGLS-C08	Gacek et al.,2021
	KenC-8 × N53-2	DH	cqGC.A9-5、cqGC.A9-7、cqGC.A9-9、 cqGC.C2-9、cqGC.C2-10、cqGC.C9-5、cqGC.C9-6	赵卫国等,2024
	19C19 × 19F105	RIL	RIL_qGS.C02	马小俊,2024
	Ken-C8 × N53-2	DH	qI3C-21-A01-1、qI3C-21-A07-1、qI3C-21-C04-1、qI3C-22-A07-1、qI3C-22-A07-2、qI3C-22-A07-3、qI3C-22-C04-1	Xiong et al.,2025
芥菜型油菜	G266 × G302	RIL	SIN-2016、SIN-2017、GNA-2016、GNA-2017	Khattak et al,2019
B. juncea L.	YufengZC × Qichi881	RIL	cQTLA02、cQTLA03、cQTLA04、cQTLB05、cQTLB06、cQTLB08	张金泽等,2025
结球甘蓝 B. oleracea L. var. capitata L.	R4-P1 × R2-P2	F₂	qGS-7-1、qGS-8-1、qGS-8-2	田多成等,2014
甘蓝B. oleracea L.	747 × 748	F₃	qSIN、q4MGBS、qGBS	Oh et al.,2018
芜菁B. napus var. napobrassica	Rutabaga-BF × UAAlfaGold	DH	qGSLs_A9	Hasan et al.,2021
白菜B. rapa L.	3-0026.027 × Sampad	RIL	GSLA2、GSLA7、GSLA9	Rahman et al.,2014
	Z16 × L144	BC₂DH	NAP-QTL3、GBN-QTL3、GBN-QTL10、PRO-QTL3、GBC-QTL9、NAS-QTL9、Total-QTL3、Ali-QTL3、Ali-QTL7、GBN-QTL5、PRO-QTL6、PRO-QTL9、GBC-QTL3、NEO-QTL9、4OH-QTL2、4OH-QTL4、NAS-QTL6、Ali-QTL5、GBN/Ali-QTL3	廖永翠,2011
	Z16 × L143	BC₂DH	NAP-QTL-A03、NAP-QTL-A09	王辉等,2011
萝卜 Raphanus sativus L.	AZ26H × TBS	F₂	GSL-QTL-1、GSL-QTL-2、GSL-QTL-3、GSL-QTL-4、GSL-QTL-5	Zou et al.,2013

物种 Species	杂交组合 Hybridized combination	群体类型 Group type	关键QTL定位 Key QTL localization	文献 Reference
甘蓝型油菜 Brassica napus L.	GH06 × P174	RIL	qGSL-A03-1、qGSL-A09-2、qGSL-A09-3、qGSL-C02-1、qGSL-A01-11、qGSL-A07-12、qGSL-A09-11、qGSL-C02-12、qGSL-A06-11	荐红举等,2014
	GH06 × 油研2号 GH06 × Youyan 2	RIL	qPRO-10-1、qPRO-14-2、qPRO-24-3、qNAP-3-1、qNAP-10-2、qNAP-24-3、qGBN-9-1、qGBN-20-2、qNAS-9-1、qNAS-15-2、qNAS-21-3	刘水燕等,2014
	EXPRESS × SWU07	DH	qGSLEA04-1、qGSLEA06-1、qGSLEA09-1、qGSLEA09-2	He et al.,2018
	G120 × 9172	DH	qGSL-C2、qGSL-A3	Liu et al.,2020b
	ZS11 × G120	DH	cqSGC-A1-1、cqSGC-A1-2、cqSGC-A1-3、cqSGC-A1-4、cqSGC-A2-1、cqSGC-A2-2、cqSGC-A3、cqSGC-A4、cqSGC-A5、cqSGC-A6-1、cqSGC-A6-2、cqSGC-A7、cqSGC-C2	周显明,2021
	M305 × Z114	DH	qGLS-A02、qGLS-A04、qSCC/GLS-C02r、 qSCC/ADF/NDF/GLS-A04、qSCC/NDF/GLS-C06、qSCC/NDF/GLS-C08、qGLS-C08	Gacek et al.,2021
	KenC-8 × N53-2	DH	cqGC.A9-5、cqGC.A9-7、cqGC.A9-9、 cqGC.C2-9、cqGC.C2-10、cqGC.C9-5、cqGC.C9-6	赵卫国等,2024
	19C19 × 19F105	RIL	RIL_qGS.C02	马小俊,2024
	Ken-C8 × N53-2	DH	qI3C-21-A01-1、qI3C-21-A07-1、qI3C-21-C04-1、qI3C-22-A07-1、qI3C-22-A07-2、qI3C-22-A07-3、qI3C-22-C04-1	Xiong et al.,2025
芥菜型油菜	G266 × G302	RIL	SIN-2016、SIN-2017、GNA-2016、GNA-2017	Khattak et al,2019
B. juncea L.	YufengZC × Qichi881	RIL	cQTLA02、cQTLA03、cQTLA04、cQTLB05、cQTLB06、cQTLB08	张金泽等,2025
结球甘蓝 B. oleracea L. var. capitata L.	R4-P1 × R2-P2	F₂	qGS-7-1、qGS-8-1、qGS-8-2	田多成等,2014
甘蓝B. oleracea L.	747 × 748	F₃	qSIN、q4MGBS、qGBS	Oh et al.,2018
芜菁B. napus var. napobrassica	Rutabaga-BF × UAAlfaGold	DH	qGSLs_A9	Hasan et al.,2021
白菜B. rapa L.	3-0026.027 × Sampad	RIL	GSLA2、GSLA7、GSLA9	Rahman et al.,2014
	Z16 × L144	BC₂DH	NAP-QTL3、GBN-QTL3、GBN-QTL10、PRO-QTL3、GBC-QTL9、NAS-QTL9、Total-QTL3、Ali-QTL3、Ali-QTL7、GBN-QTL5、PRO-QTL6、PRO-QTL9、GBC-QTL3、NEO-QTL9、4OH-QTL2、4OH-QTL4、NAS-QTL6、Ali-QTL5、GBN/Ali-QTL3	廖永翠,2011
	Z16 × L143	BC₂DH	NAP-QTL-A03、NAP-QTL-A09	王辉等,2011
萝卜 Raphanus sativus L.	AZ26H × TBS	F₂	GSL-QTL-1、GSL-QTL-2、GSL-QTL-3、GSL-QTL-4、GSL-QTL-5	Zou et al.,2013

物种 Species	材料数量 Material quantity	关联的QTL或关键基因 Associated QTL or key genes	染色体 Chromosome	文献 Reference
甘蓝型油菜 Brassica napus L.	472	BnaA.HAG1c、BnaC.HAG1a、BnaC.HAG1b、BnaC.HAG1c	A9、C2、C7、C9	Li et al.,2014
甘蓝型油菜 Brassica napus L.	520	GTR2、MYB34、MYB28	A09、C09	Qu et al.,2015
	157	MYB28、MAM1、MYB34	A09、C09、C02	魏大勇等,2018
	189	APK1、AOP1、BAT5、BGLU34、BGLU37、BGLU38、GTR1、GTR2、IGMT1、IGMT4、MAM1、MYB28、MYB34、AOP3、MYB118、NSP1	A2、A5、A6、A9、C1、C2、C9	Wang et al.,2018
	307	Homeodomain-like superfamily protein、CDPK9、RING/U-box superfamily protein、Mitochondrial transcription termination factor family protein、WD-40 repeat family protein、AOP3、MYB28	A09、C02、C09	Wei et al,2019
	288	Bna.HAG1.A9、Bna.HAG1.C2、Bna.HAG3.A3	A9、C2、A3	Kittipol et al.,2019
	366	GTR2、MYB34、GSL-OH、AOP1、BCAT4、MAM1、MYB28、CYP83A1、UGT74C1、MYB118、IGMT5、ST5b、APK2、FMO-GSOX5、IQD1、CYP81F4、CYP81F1、GSTU20、BAT5	A2、A3、A4、A5、A6、A7、A8、A9、C1、C2、C3、C4、C6、C7、C9	Liu et al.,2020a
	209	MYB28、MYB34、BCAT6、CYP71A13、MYC2、MYB122、SOT16、FMO_GS-OX1	A02、A03、A08、A09、C01、C02、C03、C09	秦晗,2021
	312	BnaA02.GTR2、BnaA09.GTR2a、BnaA09.GTR2b、BnaA09.GTR2c、BnaC02.GTR2	A02、A09、C02	He et al.,2022
	505	qGSL.A09.2、qGSL.C09.1、qGSL.C02.1、qGSL.A02.2、qGSL.A08.1、qGSL.C09.2、qGSL.A09.1、qGSL.C07.2、qGSL.C08.1、qGSL.A09.4、qGSL.C07.1、qGSL.C09.3、qGSL.C03.1、qGSL.A09.3、qGSL.A02.1	A09、C09、C02、A02、A08、C07、C08、C03	Tan et al,2022
	300	APS1、MYB34、NPF、DXR、MYB28、APK1	A05、A09、C09	周会汶等,2024
芥菜型油菜 B. juncea L.	92	HB16、SK1、CM1、JMT、LINC、CYP81G1、MYB44	A04、A06、B03、B04、B06	Akhatar et al.,2020
芥菜 B. juncea L.	183	MYB28-A02、MYB28-A09、MAM1-A02	A02、A09	李樟萍,2022
	158	GSTF11、SCPL17、OBP2、AAP4、SAL1、CYP18-3、2-ODD、MYB28、MYB34、FMOGS-OX5、PSAT1、LSU2、GRXS11、UGT71C3、RER3、CYSD1、HY5、SDI2、ALDH2B7	A02、B02、B08、A03、B04、A10、B01、B03	Tandayu et al.,2022
萝卜 R. sativus L.	179	RsGSL-OH1、RsGSL-OH2、RsGSL-OH3、Rs2OGD	Chr3	范莲雪,2020
白菜 B. rapa L.	195	IMD1、TT4、MYB119、ILR3、IQD1、PYK10、WRKY17、ADS2、NIG1、SOT18	A02、A03、A04、A06、A09	Kaur et al.,2021
	159	MYB34、MYB51、MYB28、MYB29、MYB122、SOT18、SOT16、WD40、FAH、MYB44/MYB77、MYB60、UGT74D1、UGT74C1、MYB77、CYP79B2、CHLI2、ALY3、CYP72A9、FVE、MYB118、AKN2、GTR1、IGMT4	A01、A02、A03、A04、A05、A06、A07、A08、A09	Shang et al.,2022

物种 Species	材料数量 Material quantity	关联的QTL或关键基因 Associated QTL or key genes	染色体 Chromosome	文献 Reference
甘蓝型油菜 Brassica napus L.	472	BnaA.HAG1c、BnaC.HAG1a、BnaC.HAG1b、BnaC.HAG1c	A9、C2、C7、C9	Li et al.,2014
甘蓝型油菜 Brassica napus L.	520	GTR2、MYB34、MYB28	A09、C09	Qu et al.,2015
	157	MYB28、MAM1、MYB34	A09、C09、C02	魏大勇等,2018
	189	APK1、AOP1、BAT5、BGLU34、BGLU37、BGLU38、GTR1、GTR2、IGMT1、IGMT4、MAM1、MYB28、MYB34、AOP3、MYB118、NSP1	A2、A5、A6、A9、C1、C2、C9	Wang et al.,2018
	307	Homeodomain-like superfamily protein、CDPK9、RING/U-box superfamily protein、Mitochondrial transcription termination factor family protein、WD-40 repeat family protein、AOP3、MYB28	A09、C02、C09	Wei et al,2019
	288	Bna.HAG1.A9、Bna.HAG1.C2、Bna.HAG3.A3	A9、C2、A3	Kittipol et al.,2019
	366	GTR2、MYB34、GSL-OH、AOP1、BCAT4、MAM1、MYB28、CYP83A1、UGT74C1、MYB118、IGMT5、ST5b、APK2、FMO-GSOX5、IQD1、CYP81F4、CYP81F1、GSTU20、BAT5	A2、A3、A4、A5、A6、A7、A8、A9、C1、C2、C3、C4、C6、C7、C9	Liu et al.,2020a
	209	MYB28、MYB34、BCAT6、CYP71A13、MYC2、MYB122、SOT16、FMO_GS-OX1	A02、A03、A08、A09、C01、C02、C03、C09	秦晗,2021
	312	BnaA02.GTR2、BnaA09.GTR2a、BnaA09.GTR2b、BnaA09.GTR2c、BnaC02.GTR2	A02、A09、C02	He et al.,2022
	505	qGSL.A09.2、qGSL.C09.1、qGSL.C02.1、qGSL.A02.2、qGSL.A08.1、qGSL.C09.2、qGSL.A09.1、qGSL.C07.2、qGSL.C08.1、qGSL.A09.4、qGSL.C07.1、qGSL.C09.3、qGSL.C03.1、qGSL.A09.3、qGSL.A02.1	A09、C09、C02、A02、A08、C07、C08、C03	Tan et al,2022
	300	APS1、MYB34、NPF、DXR、MYB28、APK1	A05、A09、C09	周会汶等,2024
芥菜型油菜 B. juncea L.	92	HB16、SK1、CM1、JMT、LINC、CYP81G1、MYB44	A04、A06、B03、B04、B06	Akhatar et al.,2020
芥菜 B. juncea L.	183	MYB28-A02、MYB28-A09、MAM1-A02	A02、A09	李樟萍,2022
	158	GSTF11、SCPL17、OBP2、AAP4、SAL1、CYP18-3、2-ODD、MYB28、MYB34、FMOGS-OX5、PSAT1、LSU2、GRXS11、UGT71C3、RER3、CYSD1、HY5、SDI2、ALDH2B7	A02、B02、B08、A03、B04、A10、B01、B03	Tandayu et al.,2022
萝卜 R. sativus L.	179	RsGSL-OH1、RsGSL-OH2、RsGSL-OH3、Rs2OGD	Chr3	范莲雪,2020
白菜 B. rapa L.	195	IMD1、TT4、MYB119、ILR3、IQD1、PYK10、WRKY17、ADS2、NIG1、SOT18	A02、A03、A04、A06、A09	Kaur et al.,2021
	159	MYB34、MYB51、MYB28、MYB29、MYB122、SOT18、SOT16、WD40、FAH、MYB44/MYB77、MYB60、UGT74D1、UGT74C1、MYB77、CYP79B2、CHLI2、ALY3、CYP72A9、FVE、MYB118、AKN2、GTR1、IGMT4	A01、A02、A03、A04、A05、A06、A07、A08、A09	Shang et al.,2022

基因家族 Gene family		基因名称 Gene name	物种 Species		硫苷变化 The changes of glucosinolates	参考文献 Reference
过表达Overexpression
MAM	BnMAM1		甘蓝型油菜 B. napus L.		叶片中的脂肪族硫苷含量增加1.5 ~ 1.7倍 The aliphatic GSL content in leaves increased 1.5-1.7-fold	Zhang et al.,2015b
MAM	BoMAM1		青花菜 B. oleracea L.		SF含量增加1.7 ~ 3.4倍 The SF content increased 1.7 - 3.4-fold	Cao et al.,2021
CYP83	BnCYP83A1		甘蓝型油菜 B. napus L.		叶片中的脂肪族硫苷含量增加1.8 ~ 2.2倍 The aliphatic GSL content in leaves increased 1.8-2.2-fold	Zhang et al.,2015b
CYP83	BoCYP83A1		青花菜 B. oleracea L. var. italica		GRA含量增加1.02 ~ 1.2倍，脂肪族硫苷含量增加2.11 ~ 2.29倍 The GRA content increased 1.02-1.2-fold，and the aliphatic glucosinolate content rose 2.11-2.29-fold	费丹丹,2023
SOT	BnSOT-like1		甘蓝型油菜 B. napus L.		种子中的硫苷含量增加7.33倍 The GSL content in seeds increased 7.33-fold	Wang et al.,2020c
UGT74	BoaUGT74B1		芥蓝 B. oleracea var. alboglabra		吲哚族硫苷含量增加1.30 ~ 7.11倍 The indolic GSL content increased 1.30-7.11-fold	Zheng et al.,2021
	BnUGT74B1		甘蓝型油菜 B. napus L.		叶片中的脂肪族硫苷和吲哚族硫苷含量分别增加1.7倍和1.5倍 The aliphatic and indolic GSL contents in leaves increased 1.7-fold and 1.5-fold，respectively	Zhang et al.,2015b
FMO_GS-OX	BoFMO_GS-OX2		青花菜 B. oleracea L.		SF含量增加1.6 ~ 2.7倍 The SF content increased 1.6-2.7-fold	Cao et al.,2021
FMO_GS-OX	BoFMO_GS-OX5.1		青花菜 B. oleracea L. var. italica		GRA含量增加 The GRA content increased	赵泳瑜,2024
AOP/ODD	BocODD1/2		芥蓝B. olearcea var. chinensis		PRO含量增加The PRO content increased	Wu et al.,2022
MYR	BoMYR		青花菜 B. oleracea L.		SF含量增加3.7倍 The SF content increased 3.7-fold	Cao et al.,2021
MYB	BoaMYB28		芥蓝 B. oleracea var. alboglabra		脂肪族硫苷含量增加1.5 ~ 3.0倍 The aliphatic GSL content increased 1.5-3.0-fold	Yin et al.,2017
	BoMYB28		青花菜 B. oleracea L. var. italica		GRA含量增加 The GRA content increased	杨庆喜,2024
	BnaC2.MYB28		甘蓝型油菜 B. napus L.		种子硫苷含量增加 The GSL content in seeds increased	Zhou et al.,2023
	BoMYB29		甘蓝 B. oleracea L.		脂肪族硫苷含量增加 The aliphatic GSL content increased	Zuluaga et al.,2019
	BoaMYB51		芥蓝 B. oleracea var. alboglabra		吲哚族硫苷含量增加 The indolic GSL content increased	王梦雨,2021
RNA干涉RNA interfere
MAM	BrrMAM		蔓菁 B. rapa var. rapa		GNA、PRO和GBN含量下降 The GNA，PRO and GBN contents decreased	Yin et al.,2023
CYP83	BoCYP83A1		青花菜 B. oleracea L. var. italica		GRA含量下降55% ~ 80%，脂肪族硫苷含量也下降 The GRA content decreased to 55%-80%，and the aliphatic GSL also decreased	费丹丹,2023
FMO_GS-OX	BoFMO_GS-OX5.2		青花菜 B. oleracea L. var. italica		GRA含量下降 The GRA content decreased	赵泳瑜,2024
AOP/ODD	BjuAOP2		芥菜型油菜 B. juncea L.		种子中的GRA含量增加，总硫苷含量下降；苗期中的GRA含量增加超100倍，脂肪族硫苷含量下降，总硫苷含量仅为野生型下降58% GRA content increased in seeds while total GSL decreased；in seedlings，GRA content rose over 100-fold，aliphatic GSL decreased，and total GSL was only 58% of the wild-type	Augustine & Bisht,2015
AOP/ODD	BnAOP2		甘蓝型油菜 B. napus L.		种子中的GRA含量增加，PRO含量下降 The GRA content in seeds increased，while the PRO content decreased	Liu et al.,2012
	BrrAOP2		蔓菁 B. rapa var. rapa		GRA含量增加 The GRA content increased	Yin et al.,2023
	BocODD1/2		芥蓝 B. oleracea var. chinensis		PRO含量下降 The PRO content decreased	Wu et al,2022
GTR	BocGTR1a		芥蓝 B. oleracea var. chinensis		叶片中的硫苷含量下降，根部中的硫苷含量增加 The GSL content in leaves decreased，while the GSL content in roots increased	Jiang et al.,2019
GTR		BjuGTR1	芥菜型油菜 B. juncea L.		种子和叶片中的硫苷含量下降 The GSL content in seeds and leaves decreased	Nambiar et al.,2021
		BjuGTR2	芥菜型油菜 B. juncea L.		种子中的硫苷含量下降，叶荚果中的硫苷含量增加 The GSL content in seeds decreased，while the GSL content in leaf pods increased	Nambiar et al.,2021
MYB		BoaMYB28	芥蓝B. oleracea var. alboglabra		脂肪族硫苷含量下降 The aliphatic GSL content decreased	Yin et al.,2017
		BjMYB28	芥菜型油菜 B. juncea		叶片和种子中的脂肪族硫苷含量下降 The content of aliphatic GSL in leaves and seeds decreased	Augustine et al.,2013
		BnaC2.MYB28	甘蓝型油菜 B. napus L.		种子中的硫苷含量下降 The GSL content in seeds decreased	Zhou et al.,2023
基因编辑Genome editing
AOP/ODD		BoaAOP2	芥蓝 B. oleracea var. alboglabra		GRA含量增加 The GRA content increased	袁巧,2021
		BoAOP2	甘蓝 B. oleracea. var. capitata		GRA含量增加，PRO含量下降 The GRA content increased，while the PRO content decreased	王鹏,2023
		BoaAOP2	芥蓝 B. oleracea L.		GRA含量增加11.71 ~ 41.29倍，脂肪族硫苷含量下降 The GRA content increased by 11.71-41.29-fold，while the aliphatic GSL content decreased	Zheng et al,2023
GTR		BnaA06.GTR2	甘蓝型油菜 B. napus L.		种子中的硫苷含量下降76.05% The GSL content in seeds decreased by 76.05%	He et al.,2022
		BnaA02.gtr2/ BnaC02.gtr2/ BnaC03.gtr2/ BnaA06.gtr2		甘蓝型油菜 B. napus L.	在单突变体种子的硫苷含量中，BnaA02.gtr2和BnaC02.gtr2的下降最为显著，BnaC03.gtr2和BnaA06.gtr2呈中度下降；双突变体和三突变体的硫苷含量较单突变体进一步下降，四突变体则下降86.85% In single-mutant seeds，GSL content decreased most significantly in BnaA02.gtr2 and BnaC02.gtr2，moderately in BnaC03.gtr2 and BnaA06.gtr2；GSL content in double and triple mutants was further reduced compared with single mutants，while that in quadruple mutants decreased by 86.85%	Tan et al.,2022
MYB		BoMYB28	甘蓝 B. oleracea L.		叶片中的GRA含量下降 The GRA content in leaves decreased	Neequaye et al.,2021
MYB		BnaC2.MYB28	甘蓝型油菜B. napus L.		种子中的硫苷含量下降The GSL content in seeds decreased	Zhou et al.,2023