上位基因控制的茄子果色遗传效应解析

doi:10.3864/j.issn.0578-1752.2023.23.014

中国农业科学 ›› 2023, Vol. 56 ›› Issue (23): 4729-4741.doi: 10.3864/j.issn.0578-1752.2023.23.014

上位基因控制的茄子果色遗传效应解析

陈锦溶¹^,²(), 吕紫见¹^,³, 范丽莎¹^,², 游倩¹, 李涛¹, 宫超¹, 孙光闻², 李植良¹(), 孙保娟¹()

¹ 广东省农业科学院蔬菜研究所/广东省蔬菜新技术研究重点实验室，广州 510640
² 华南农业大学园艺学院，广州 510642
³ 华中农业大学园艺植物生物学教育部重点实验室，武汉 430070

收稿日期:2023-06-05 接受日期:2023-10-04 出版日期:2023-12-04 发布日期:2023-12-04
通信作者:
孙保娟，Tel：020-38469456；E-mail：sunbaojuan@hotmail.com。
李植良，E-mail：vri_li@163.com
联系方式: 陈锦溶，E-mail：20213137106@stu.scau.edu.cn。
基金资助:
广西科技重大专项(桂科AA22068088); 广东省现代农业产业技术体系(2023KJ110); 广东省现代农业产业技术体系(2023KJ106); 广东省级乡村振兴战略专项资金种业振兴项目(2022-NPY-00-026)

Analysis of Genetic Effect of Fruit Color Controlled by Epistatic Genes in Eggplant

CHEN JinRong¹^,²(), LÜ ZiJian¹^,³, FAN LiSha¹^,², YOU Qian¹, LI Tao¹, GONG Chao¹, SUN GuangWen², LI ZhiLiang¹(), SUN BaoJuan¹()

¹ Vegetable Research Institute, Guangdong Academy of Agricultural Sciences/Guangdong Key Laboratory for New Technology Research of Vegetables, Guangzhou 510640
² College of Horticulture, South China Agricultural University, Guangzhou 510642
³ Key Laboratory of Horticultural Plant Biology, Ministry of Education, Huazhong Agricultural University, Wuhan 430070

Received:2023-06-05 Accepted:2023-10-04 Published:2023-12-04 Online:2023-12-04

摘要/Abstract

摘要：

【目的】果实颜色是影响茄子商品价值的重要性状。通过解析两白果亲本杂交构建的F₂代群体紫红果单株和白果单株特殊分离比产生的原因，为阐明茄子果实颜色形成的调控机制奠定基础。【方法】以花青素合成结构基因ANS（P）突变的白花白果母本19141、未知突变基因位点的白花白果父本19142及其紫红果F₁、果色分离的F₂群体为试验材料，探讨上位基因控制的茄子果色遗传规律；克隆已知的茄子果色相关D（SmMYB1）和Y（SmDFR）基因，探明父本19142突变的果色基因和方式；开发基因内分子标记，对F₂进行基因分型以及与其他果皮无花青素沉着茄子亲本杂交验证，解析上位基因调控茄子果色遗传的分子基础。【结果】E4450 F₂紫红果和白果单株分离比符合3对上位基因控制的27﹕37的分离比率，19141的P基因位点发生突变，基因型为DDppYY，19142的D和Y基因位点同时发生了突变，基因型为ddPPyy。克隆测序发现，19142的SmMYB1发生可变剪接，导致第2外显子跳跃。19142的SmDFR起始密码子上游-326 bp的SNP（C→G），导致启动子缺失了1个CAAT-box顺式作用元件；在第2外显子最后一个碱基G变C，发生剪切位点突变。基于SmMYB1、SmANS和SmDFR遗传变异，开发了基因内分子标记对E4450 F₂单株进行基因分型，结果发现基因型与表型完全吻合；D_P_Y_对应紫花紫红果表型，ddP_Y_基因型对应紫花白果表型，D_ppY_、D_P_yy、D_ppyy、ddppY_、ddP_yy和ddppyy基因型对应白花白果表型。19142与白果自交系19147（d^td^tPPYY）和绿果自交系19144（DDPPyy）分别杂交，结果发现F₁果色分别为白色和绿色，果皮无花青素沉着，这进一步证明了19142是SmMYB1和SmDFR两个基因同时发生突变。【结论】2个果皮无花青素沉着的茄子亲本杂交，如果F₁代果皮有花青素沉着，且F₂代有花青素沉着单株和无花青素沉着果色单株分离比符合27﹕37，则是由于其中的一个亲本在花青素合成通路的1个基因位点发生了突变，另外一个亲本在花青素合成通路的另外2个基因位点发生了突变。两个果皮无花青素沉着亲本杂交，F₁果皮能够合成花青素而呈现紫红，是由于3个上位基因位点D、P和Y同时处于显性状态，花青素合成得以恢复。结构基因SmANS或SmDFR突变抑制植株各个部位花青素合成；转录因子SmMYB1调控具有组织特异性，其突变抑制果皮花青素合成，却不抑制花中花青素合成。

关键词: 茄子, 果色, 上位基因, 花青素, 遗传规律

Abstract:

【Objective】Fruit color is an important trait that affects the commercial value of eggplant fruit. By analyzing the causes for the special segregation ratio of individual plants with purple red peel and with white peel in the F₂ population constructed by crossing between two white-fruit parents, this paper could lay the foundation for elucidating the mechanism of epistatic gene interaction on regulating eggplant fruit coloration.【Method】The white-flower and white-peel female parent 19141 with mutation at the structural gene ANS involving in the anthocyanin biosynthesis pathway, white-flower and white-peel male parent 19142 with unknown mutation genes involving in the anthocyanin biosynthesis pathway, and their F₁ population with purple red peel and F₂population with separate peel colors were used to explore the epistatic inheritance of eggplant fruit coloration. Genes and their mutation patterns were studied by cloning the known genes, D (SmMYB1) and Y (SmDFR) related to peel color in male parent 19142. Molecular basis of peel color-controlling epigenetic genes was analyzed by developing molecular markers based on genetic variations of peel color genes, analyzing the relationship between genotype and phenotype in E4450F₂ population, and crossing with other eggplant parents without anthocyanin pigmentation in peel.【Result】The segregation ratio of plants with purple red fruit and white fruit in E4450 F₂ progeny was consistent with the segregation ratio of 27:37 controlled by three pairs of epistatic genes, that is, mutations occurred at the P gene locus in 19141, with genotype DDppYY, and mutations occurred at both D and Y gene loci in 19142, with genotype ddPPyy. The results of cloning and sequencing showed that alternative splicing occurred in SmMYB1 of 19142, which led to the second exon skipping. In 19142, SNP (C→G) in the promoter region’s -326 bp upstream of the start codon resulted in the absence of a CAAT-box cis-acting element in SmDFR gene. An SNP, G to C, at the last base of the second exon, was annotated as splicing mutation, which might cause abnormal function of SmDFR gene in 19142, resulting in the eggplant peel’s inability to synthesize anthocyanin. Based on the genetic variation of SmMYB1, SmANS and SmDFR, the functional molecular markers were developed, and the progenies of E4450F₂ were genotyped. The results showed that genotype and phenotype were completely consistent. D_P_Y_ corresponded to phenotypes of purple flower and purple red peel, ddP_Y_ corresponded to phenotype of purple flower and white peel, D_ppY_, D_P_yy, D_ppyy, ddppY_, ddP_yy and ddppyy genotypes corresponded to white flower and white peel phenotypes. When 19142 was crossed with white-peel inbred line 19147 (d^td^tPPYY), and green-peel inbred line 19144 (DDPPyy), it was found that the fruit color of the two F₁progenies were white and green, respectively, and there was no anthocyanin pigmentation in the peel, which further proved that 19142 was a double mutant in SmMYB1 and SmDFR.【Conclusion】When two eggplant parents without anthocyanin pigmentation in the peel were crossed, the peel of the F₁ generation had anthocyanidin pigmentation, and the segregation ratio of plants with anthocyanin pigmentation and non-anthocyanin pigmentation in F₂ population was 27:37, it was because one of the parents had a mutation at a gene locus in the anthocyanin biosynthesis pathway, and the other parent had mutations at two other loci in the anthocyanin biosynthesis pathway. When two parents without anthocyanin pigmentation were crossed, the peel of F₁ was able to synthesize anthocyanin and present purple red color, which was due to the simultaneous dominance of three epistatic gene loci D, P and Y, and anthocyanin biosynthesis was restored. Mutation of the structural gene of SmANS or SmDFR inhibited anthocyanin biosynthesis in all parts of the plant. The regulation of transcription factor mutation SmMYB1 was tissue specific, and its mutation inhibited anthocyanin biosynthesis in the peel, but did not inhabit anthocyanin biosynthesis in the flower.

Key words: eggplant, peel color, epistatic gene, anthocyanin, genetic law

陈锦溶, 吕紫见, 范丽莎, 游倩, 李涛, 宫超, 孙光闻, 李植良, 孙保娟. 上位基因控制的茄子果色遗传效应解析[J]. 中国农业科学, 2023, 56(23): 4729-4741.

CHEN JinRong, LÜ ZiJian, FAN LiSha, YOU Qian, LI Tao, GONG Chao, SUN GuangWen, LI ZhiLiang, SUN BaoJuan. Analysis of Genetic Effect of Fruit Color Controlled by Epistatic Genes in Eggplant[J]. Scientia Agricultura Sinica, 2023, 56(23): 4729-4741.

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用途 Usage	引物名称 Primer name	引物序列 Primer sequence (5′-3′)	退火温度 Anneal temperature (℃)	延伸时间 Extending time (s)
克隆SmANS全长 Cloning of the full length SmANS	ANS-B1	F: CCAGTCGTTTGTTCCCACAT R: GGACATTACCAGAGGGTTGA	55	30
	ANS-B2	F: TACCTTCTTTGATCTGCCTGTT R: GCTCTTGTACTTTCCGTTGC	60	60
	ANS-B3	F: TCATCCTCCACAATATGGTGC R: CTGTGTTGCTCCTTCACAAAGG	60	30
克隆SmMYB1全长 Cloning of the full length SmMYB1	MYB1-D1	F: TCCGTACAAGGTGGCAGGATAGGTTATTAGG R: CGGAAGTCTACCAGCAATAAGTGACCATCTA	60	60
克隆SmMYB1全长 Cloning of the full length SmMYB1	MYB1-C2	F: GTGCAATGATGACGCTCACA R: GGACCTATGCCGAAAACAGA	55	30
克隆SmMYB1 cDNA Cloning of cDNA of SmMYB1	MYB1-CDS1	F: ATGAATAATCCTCCTATAATCTGTACGTC R: CTGCTCTAGCAGGAACAAGAT	53	30
	MYB1-CDS2	F: TAATCTGTACGTCTGTGCGAGTGA R: GTTATTTCTTTGCAGCGTTCTTCC	55	30
	MYB1-CDS3	F: TGGAACACTAACCTTCTAAGG R: TTAAACAAGTAGATTCCATAAATCAAT	50	30
克隆SmDFR全长 Cloning of the full length SmDFR	65-1036	F: AGGTCTCCCAAACCCACAAC R: CGCTCCAGCTAGTCTCGTCA	55	60
	DFR-A2	F: CTGTCAGGGAGTATTTCATGTG R: AGCTTCTTCCCTTGCAGC	55	30
	1060-1900	F: AAGATGACAGGATGGGTTCG R: CCTTGCCACTTGCATAGTTT	53	60
	DFR-A4	F: GTATACTTGGCTCTTTCATAGAGTG R: CTGCCTGAAATTAAGCAAGATGAG	53	30
克隆SmDFR启动子 Cloning of the promoter of SmDFR	DP2	F: GGCTTCCTTCGTAGTCAA R: CCCGCAAGTCTAATCTCAC	53	60
	3-403	F: CCGTAACAAATATCAGCATTTC R: GAGAGAATCGTATTTTTGAAAGC	50	30
	DFR-qA4	F: CTGATGTTCATGGATAGACGGA R: TGCTTGTACGTACATGATGTTAGG	53	30

标记 Marker	引物名称 Primer name	引物序列 Primer Sequence (5′-3′)
SmMYB1-26bpInDel	SmMYB1 FF2	TATAATCTGTACGTCTGTGCGAGTG
SmMYB1-26bpInDel	SmMYB1 FR2	GCCTTAAATAATTCAACCACCTCA
SmMYB1-52bpInDel	MYB1-52FF	TAGGCTGAGGTGGTTGAAT
SmMYB1-52bpInDel	MYB1-52FR	TTCGCTAAATATATTATTGTGGAGAC
SBJ-DFR	SBJ-DFR-Fg	GAAGGTGACCAAGTTCATGCTTCGAGTCCAAGGACCCTGAG
	SBJ-DFR-Fc	GAAGGTCGGAGTCAACGGATTTCGAGTCCAAGGACCCTGAC
	SBJ-DFR-R	AGGCAATAAGTATAGATGAGTTAACTCTG
SmANS-Chr08SNP	SmANS-Chr08SNP-F	CCTAAAGAGTATGTGAGGCCACAAG
	SmANS-Chr08SNP-Rt	GAAGGTGACCAAGTTCATGCTCCTCAAAGATGTTTCCTATACCATTTT
	SmANS-Chr08SNP-Ra	GAAGGTCGGAGTCAACGGATTCCTCAAAGATGTTTCCTATACCATTTA

群体 Population	总株数 Total plants	紫红果单株数 Number of purple red peel plants Anthocyanin pigmented	白果单株数 Number of white peel plants	卡方检验x² (27﹕37)	P值 P value
19141	30	-	30	-	-
19142	30	-	30	-	-
E4450F₁后代 E4450F₁ descendants	30	30	0	-	-
E4450F₂群体 E4450F₂ population	247	106	141	0.054	0.817

	F₁的配子 Gametes of F₁
	DPY	DpY	DPy	Dpy	dPY	dpY	dPy	dpy
DPY	DDPPYY	DDPpYY	DDPPYy	DDPpYy	DdPPYY	DdPpYY	DdPPYy	DdPpYy
DpY	DDPpYY	DDppYY	DDPpYy	DDppYy	DdPpYY	DdppYY	DdPpYy	DdppYy
DPy	DDPPYy	DDPpYy	DDPPyy	DDPpyy	DdPPYy	DdPpYy	DdPPyy	DdPpyy
Dpy	DDPpYy	DDppYy	DDPpyy	DDppyy	DdPpYy	DdppYy	DdPpyy	Ddppyy
dPY	DdPPYY	DdPpYY	DdPPYy	DdPpYy	ddPPYY	ddPpYY	ddPPYy	ddPpYy
dpY	DdPpYY	DdppYY	DdPpYy	DdppYy	ddPpYY	ddppYY	ddPpYy	ddppYy
dPy	DdPPYy	DdPpYy	DdPPyy	DdPpyy	ddPPYy	ddPpYy	ddPPyy	ddPpyy
dpy	DdPpYy	DdppYy	DdPpyy	Ddppyy	ddPpYy	ddppYy	ddPpyy	ddppyy

果色 Peel color	花色 Flower color	SmMYB1-26bpInDel的基因型 Genotype of SmMYB1-26bpInDel	SmANS-Chr08SNP的基因型 <BOLD>G</BOLD>enotype of SmANS-Chr08SNP	SBJ-DFR的基因型 <BOLD>G</BOLD>enotype of SBJ-DFR	基因型 Genotype	株数 Plant number
紫红色Purplish red	紫色 Purple	DD	PP	YY	DDPPYY	3
紫红色Purplish red	紫色 Purple	DD	Pp	YY	DDPpYY	2
紫红色Purplish red	紫色 Purple	DD	PP	Yy	DDPPYy	2
紫红色Purplish red	紫色 Purple	DD	Pp	Yy	DDPpYy	3
紫红色Purplish red	紫色 Purple	Dd	PP	YY	DdPPYY	4
紫红色Purplish red	紫色 Purple	Dd	Pp	YY	DdPpYY	9
紫红色Purplish red	紫色 Purple	Dd	PP	Yy	DdPPYy	8
紫红色Purplish red	紫色 Purple	Dd	Pp	Yy	DdPpYy	8
白色White	白色 White	DD	pp	YY	DDppYY	2
白色White	白色 White	DD	pp	yy	DDppyy	1
白色White	白色 White	DD	pp	Yy	DDppYy	4
白色White	白色 White	DD	PP	yy	DDPPyy	2
白色White	白色 White	DD	Pp	yy	DDPpyy	1
白色White	白色 White	Dd	pp	Yy	DdppYy	8
白色White	白色 White	Dd	pp	YY	DdppYY	2
白色White	白色 White	Dd	pp	yy	Ddppyy	6
白色White	白色 White	Dd	Pp	yy	DdPpyy	6
白色White	白色 White	Dd	PP	yy	DdPPyy	4
白色White	白色 White	dd	pp	YY	ddppYY	0
白色White	白色 White	dd	pp	Yy	ddppYy	6
白色White	白色 White	dd	PP	yy	ddPPyy	1
白色White	白色 White	dd	pp	yy	ddppyy	0
白色White	白色 White	dd	Pp	yy	ddPpyy	5
白色White	紫色 Purple	dd	PP	Yy	ddPPYy	3
白色White	紫色 Purple	dd	Pp	Yy	ddPpYy	7
白色White	紫色 Purple	dd	PP	YY	ddPPYY	3
白色White	紫色 Purple	dd	Pp	YY	ddPpYY	2

上位基因控制的茄子果色遗传效应解析

Analysis of Genetic Effect of Fruit Color Controlled by Epistatic Genes in Eggplant

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