辣椒素类物质生物合成研究进展

doi:10.16420/j.issn.0513-353x.2024-0786

摘要/Abstract

摘要：

辣椒素类物质是辣椒中特异性合成的一类物质，在食品、医药、农药等领域都存在着广泛的用途。近年来很多学者对辣椒素类物质的生物合成及相关的酶和基因进行了研究，并且取得了重要进展。本文中综述了近年来辣椒素类物质的合成相关的研究，包括辣椒素类物质的种类、生物合成、合成相关的基因、含量相关的QTL以及影响其生物合成的外界环境条件等方面。

关键词: 辣椒素类物质, 基因, 生物合成, 进展

Abstract:

Capsaicinoids are a type of substance specifically synthesized in chili peppers，which is widely used in food，medicine，pesticide and other fields. In recent years，many scholars have studied the biosynthesis of capsaicinoids and related genes and enzymes，and have made important progress. In this paper，the synthesis of capsaicinoids in recent years were reviewed，including the classification of peppers，the types of capsaicinoids，the biosynthesis and genes related of capsaicinoids，QTLs related to capsaicinoids content，and the external environmental conditions affecting the capsaicinoids biosynthesis.

Key words: capsaicinoids, gene, biosynthesis, progress

温馨, 张正海, 于海龙, 吴华茂, 黄俊轩, 曹亚从, 王立浩. 辣椒素类物质生物合成研究进展[J]. 园艺学报, 2025, 52(11): 3103-3120.

WEN Xin, ZHANG Zhenghai, YU Hailong, WU Huamao, HUANG Junxuan, CAO Yacong, WANG Lihao. Research Progress on Biosynthesis of Capsaicinoids[J]. Acta Horticulturae Sinica, 2025, 52(11): 3103-3120.

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Pun1	Pun1突变位点 The mutation site of Pun1	代表材料 Representative material	参考文献 Reference
Pun1¹	具有跨越1.8 kb推定启动子和0.7 kb短截的的第一个外显子的大的2.5 kb的缺失 A large 2.5 kb deletion spanning a putative 1.8 kb promoter region and a 0.7 kb segment of 1rst exon	Maor，Jupiter，Sweet 3575 （C. annuum）	Stewart et al.,2005
Pun1²	第一个外显子区域具有4 bp的缺失 1st exon region has a 4 bp deletion	NMCA 30036 （C. chinense）	Stewart et al.,2007
Pun1³	末端70个氨基酸的缺失，提前产生终止密码子，导致了第二个外显子被短截 The deletion of 70 amino acids at the end of the second exon leads to the premature generation of a stop codon, resulting in the truncation of 2nd exon	PI594141-np （C. frutescens）	Stellari et al.,2010
Pun1⁴	第二个外显子中插入了单个核苷酸导致产生了移码突变 The insertion of a single nucleotide in 2nd exon results in a frameshift mutation	Nara Murasaki （C. annuum）	Kirii et al.,2017
Pun1⁵	包含Pun1的18.5-kb缺失 An 18.5kb deletion encompassing Pun1	Sampo Oamanaga（C. annuum）	Yamaguchi et al.,2024

Pun1	Pun1突变位点 The mutation site of Pun1	代表材料 Representative material	参考文献 Reference
Pun1¹	具有跨越1.8 kb推定启动子和0.7 kb短截的的第一个外显子的大的2.5 kb的缺失 A large 2.5 kb deletion spanning a putative 1.8 kb promoter region and a 0.7 kb segment of 1rst exon	Maor，Jupiter，Sweet 3575 （C. annuum）	Stewart et al.,2005
Pun1²	第一个外显子区域具有4 bp的缺失 1st exon region has a 4 bp deletion	NMCA 30036 （C. chinense）	Stewart et al.,2007
Pun1³	末端70个氨基酸的缺失，提前产生终止密码子，导致了第二个外显子被短截 The deletion of 70 amino acids at the end of the second exon leads to the premature generation of a stop codon, resulting in the truncation of 2nd exon	PI594141-np （C. frutescens）	Stellari et al.,2010
Pun1⁴	第二个外显子中插入了单个核苷酸导致产生了移码突变 The insertion of a single nucleotide in 2nd exon results in a frameshift mutation	Nara Murasaki （C. annuum）	Kirii et al.,2017
Pun1⁵	包含Pun1的18.5-kb缺失 An 18.5kb deletion encompassing Pun1	Sampo Oamanaga（C. annuum）	Yamaguchi et al.,2024

pAMT		pAMT突变位点 The mutation site of pAMT	代表材料 Representative material	参考文献 Reference
已命名Named	pamt¹	在第16个外显子插入1个T T insertion in 16th exon	CH-19 Sweet （C. annuum)	Lang et al.,2009
	pamt²	第11个外显子有1个核苷酸变异导致PLP结合域发生氨基酸替换 A nucleotide change in 11th exon resulted in institution of amino acid in PLP binding domain	Himo （C. annuum）	Tanaka et al.,2010a
	pamt³	第8个外显子中插入5 bp（TGGGC） 5 bp（TGGGC）insertion in 8th exon	Belize Sweet （C. chinense）	Tanaka et al.,2010b
	pamt⁴	第5个内含子插入了2.3 kb的转座子 2.3 kb insertion in 5th intron	Zavory Hot （C. chinense）	Tanaka et al.,2010b
	pamt⁵	第3个内含子插入了2.3 kb的转座子，第6个外显子插入8 bp（GCCACACC） 2.3 kb insertion in 3rd intron，8 bp（GCCACACC）insertion in 6th exon	Aji Dulce strain 2 （C. chinense）	Tanaka et al.,2010b
	pamt⁶	第2个外显子插入了7 bp（CTTTACT） 7 bp（CTTTACT）insertion in 2nd exon	No. 80（C. chinense）	Koeda et al.,2014
	pamt⁷	第2个外显子插入2.8 kb转座子 2.8 kb insertion in 2nd exon	LP6（C. chinense）	Tanaka et al.,2015
	pamt⁸	第7个外显子缺失 12 bp（TCTGCTGGTCTC） 12 bp（TCTGCTGGTCTC）deletion mutation in 7th exon	S3212（C. frutescens）	Park et al.,2015
	pamt⁹	第16个外显子中插入7 bp（TCGGTAC） 7 bp（TCGGTAC）insertion in 16th exon	No. 4034（C. chinense）	Tanaka et al.,2018
	pamt¹⁰	第11个外显子上有一个无义取代 There is a nonsense substitution in the 11th exon	Color Piman Yellow（C. annuum）	Tsurumaki & Sasanuma,2019
	pamt^L1	在内含子3中具有Tcc转座子的插入，改变了pAMT蛋白的剪接效率进而使转录水平降低，导致不同的pAMT活性 An insertion of the Tcc transposon in the third intron alters the splicing efficiency of the pAMT protein，thereby reducing its transcriptional level and resulting in varying pAMT activity	Devil’s Yellow（C. chinense）	Tanaka et al.,2019
	pamt^L2	在内含子3中具有Tcc转座子的插入，改变了pAMT蛋白的剪接效率进而使转录水平降低，导致不同的pAMT活性 An insertion of the Tcc transposon in the third intron alters the splicing efficiency of the pAMT protein，thereby reducing its transcriptional level and resulting in varying pAMT activity	LP13（C. chinense）	Tanaka et al.,2019
未命名 Unnamed	隐性等位基因Recessive allele	在第3个和第6个外显子上分别有403 bp和8 bp的插入缺失，导致产生了一个早期的终止密码子，而产生辣椒素酯类物质 An early stop codon in pAMT resulted from 403 bp and 8 bp insertions in 3rd and 6th exons	SNU11-001 （C. chinense）	Jang et al.,2015
未命名 Unnamed	pAMT（Pun2）的隐性等位基因Recessive allele of pAMT（Pun2）	第8个外显子上有一个7 bp的核苷酸插入，导致早期的终止密码子，导致产生截短的蛋白 There is a 7 bp nucleotide insertion in 8th exon，leading to an early stop codon and resulting in the production of a truncated protein	PI260433-np （C. chacoense）	Yi et al.,2022

pAMT		pAMT突变位点 The mutation site of pAMT	代表材料 Representative material	参考文献 Reference
已命名Named	pamt¹	在第16个外显子插入1个T T insertion in 16th exon	CH-19 Sweet （C. annuum)	Lang et al.,2009
	pamt²	第11个外显子有1个核苷酸变异导致PLP结合域发生氨基酸替换 A nucleotide change in 11th exon resulted in institution of amino acid in PLP binding domain	Himo （C. annuum）	Tanaka et al.,2010a
	pamt³	第8个外显子中插入5 bp（TGGGC） 5 bp（TGGGC）insertion in 8th exon	Belize Sweet （C. chinense）	Tanaka et al.,2010b
	pamt⁴	第5个内含子插入了2.3 kb的转座子 2.3 kb insertion in 5th intron	Zavory Hot （C. chinense）	Tanaka et al.,2010b
	pamt⁵	第3个内含子插入了2.3 kb的转座子，第6个外显子插入8 bp（GCCACACC） 2.3 kb insertion in 3rd intron，8 bp（GCCACACC）insertion in 6th exon	Aji Dulce strain 2 （C. chinense）	Tanaka et al.,2010b
	pamt⁶	第2个外显子插入了7 bp（CTTTACT） 7 bp（CTTTACT）insertion in 2nd exon	No. 80（C. chinense）	Koeda et al.,2014
	pamt⁷	第2个外显子插入2.8 kb转座子 2.8 kb insertion in 2nd exon	LP6（C. chinense）	Tanaka et al.,2015
	pamt⁸	第7个外显子缺失 12 bp（TCTGCTGGTCTC） 12 bp（TCTGCTGGTCTC）deletion mutation in 7th exon	S3212（C. frutescens）	Park et al.,2015
	pamt⁹	第16个外显子中插入7 bp（TCGGTAC） 7 bp（TCGGTAC）insertion in 16th exon	No. 4034（C. chinense）	Tanaka et al.,2018
	pamt¹⁰	第11个外显子上有一个无义取代 There is a nonsense substitution in the 11th exon	Color Piman Yellow（C. annuum）	Tsurumaki & Sasanuma,2019
	pamt^L1	在内含子3中具有Tcc转座子的插入，改变了pAMT蛋白的剪接效率进而使转录水平降低，导致不同的pAMT活性 An insertion of the Tcc transposon in the third intron alters the splicing efficiency of the pAMT protein，thereby reducing its transcriptional level and resulting in varying pAMT activity	Devil’s Yellow（C. chinense）	Tanaka et al.,2019
	pamt^L2	在内含子3中具有Tcc转座子的插入，改变了pAMT蛋白的剪接效率进而使转录水平降低，导致不同的pAMT活性 An insertion of the Tcc transposon in the third intron alters the splicing efficiency of the pAMT protein，thereby reducing its transcriptional level and resulting in varying pAMT activity	LP13（C. chinense）	Tanaka et al.,2019
未命名 Unnamed	隐性等位基因Recessive allele	在第3个和第6个外显子上分别有403 bp和8 bp的插入缺失，导致产生了一个早期的终止密码子，而产生辣椒素酯类物质 An early stop codon in pAMT resulted from 403 bp and 8 bp insertions in 3rd and 6th exons	SNU11-001 （C. chinense）	Jang et al.,2015
未命名 Unnamed	pAMT（Pun2）的隐性等位基因Recessive allele of pAMT（Pun2）	第8个外显子上有一个7 bp的核苷酸插入，导致早期的终止密码子，导致产生截短的蛋白 There is a 7 bp nucleotide insertion in 8th exon，leading to an early stop codon and resulting in the production of a truncated protein	PI260433-np （C. chacoense）	Yi et al.,2022

QTL名称 Name of QTL	共定位基因 Co-localized gene	染色体 Chromosome	参考文献 Reference
TH-total1.1，TH-cap1.1		Chrom01	Han et al.,2018
TH-cap1.2，TH-total1.2		Chrom01	Han et al.,2018
TH-cap1.3		Chrom01	Han et al.,2018
PD-dicap1.1，PD-cap1，PD-total1.1，TH-cap1.4		Chrom01	Han et al.,2018
PD-dicap1.2，PD-dicap1.3，PD-total1.2		Chrom01	Han et al.,2018
TH-cap1.5，TH-total1.3		Chrom01	Han et al.,2018
TH-cap2.1		Chrom02	Han et al.,2018
TH-total2，TH-cap2.2，PD-dicap2.1，PD-dicap2.2，PD-total2，PD-cap2，qdhc2.1，qdhc2.2，QTLsFP-P2.1	Pun1（CaAT3）	Chrom02	Lee et al.,2016; Han et al.,2018
3.1		Chrom03	Yarnes et al.,2013
HJ-dhc3	pAMT	Chrom03	Park et al.,2019
TH-dicap3.1，TH-dicap3.2，TH-total3.1		Chrom03	Han et al.,2018
TH-dicap3.3		Chrom03	Han et al.,2018
TH-total3.2，PD-cap3	CSE（CA03g24780）	Chrom03	Han et al.,2018
qcap3.1	CSE（CA03g24780）	Chrom03	Lee et al.,2016
Shql3		Chrom03	Kondo et al.,2023
TH-total3.3	3A2，4CL（CA03g30500）	Chrom03	Han et al.,2018
cap3.1，total3.1	3A2，4CL	Chrom03	Ben-Chaim et al.,2006
QTLFP-P3.3		Chrom03	McLeod et al.,2023
4.2		Chrom04	Yarnes et al.,2013
PD-total4.1，PD-dicap4		Chrom04	Han et al.,2018
TH-cap4，PD-total4.2		Chrom04	Han et al.,2018
cap4.1，dhc4.1，total4.1，4.13，4.14，4.15，4.16	BCAT	Chrom04	Ben-Chaim et al.,2006；Yarnes et al.,2013
5.4		Chrom05	Yarnes et al.,2013
SJ-dhc6.1		Chrom06	Park et al.,2019
HJ-tcp6.1，HJ-tcp6.2		Chrom06	Park et al.,2019
qcap6.1	IPMS（CA06g08090）	Chrom06	Lee et al.,2016
PD-cap6		Chrom06	Han et al.,2018
SJ-dhc6.2，punv	Pun4，Capana06g001204	Chrom06	Park et al.,2019；Cao et al.,2022
HJ-tcp6.4（2017）		Chrom06	Park et al.,2019
HJ-tcp6.3		Chrom06	Park et al.,2019
HJ-dhc6.1，HJ-tcp6.4（2016）		Chrom06	Park et al.,2019
HJ-dhc6.2		Chrom06	Park et al.,2019
SJ-cap6.1，SJ-dhc6.3，SJ-tcp6.1，SJ-cap6.2，SJ-tcp6.2		Chrom06	Park et al.,2019
6.8，TH-dicap6，TH-total6	FatA（CA06g26640）	Chrom06	Yarnes et al.,2013；Han et al.,2018
cap7.1，dhc7.1，total7.1		Chrom07	Ben-Chaim et al.,2006
QTLFP-P7.3		Chrom07	McLeod et al.,2023
7.3	Pun3（CaMYB31）	Chrom07	Yarnes et al.,2013
cap，cap7.2，dhc7.2，total7.2	Pun3（CaMYB31）	Chrom07	Blum et al.,2003；Ben-Chaim et al.,2006
Shql7		Chrom07	Kondo et al.,2023
PD-dicap10.1，PD-total10.1		Chrom10	Han et al.,2018
TH-total10，TH-dicap10，PD-cap10，PD-total10.2，PD-dicap10.2		Chrom10	Han et al.,2018
10.2		Chrom10	Yarnes et al.,2013
10.3	CaKR1（CA10g18840）	Chrom10	Yarnes et al.,2013
HJ-tcp11		Chrom11	Park et al.,2019
11.8		Chrom11	Yarnes et al.,2013
PD-dicap12，PD-total12		Chrom12	Han et al.,2018