万寿菊舌状花花冠裂片突变体的形态鉴定及连锁标记开发

doi:10.11983/CBB22231

植物学报 ›› 2023, Vol. 58 ›› Issue (6): 893-904.DOI: 10.11983/CBB22231

万寿菊舌状花花冠裂片突变体的形态鉴定及连锁标记开发

王文静¹^,^†, 朱钰¹^,^†, 张洪铭¹^,², 韦陆丹¹, 易庆平³, 余晓敏¹, 刘雨菡¹, 张莉雪¹, 程文翰³, 何燕红¹^,^*()

¹华中农业大学园艺林学学院, 果蔬园艺作物种质创新与利用全国重点实验室, 武汉 430070
²南京农业大学园艺学院, 作物遗传与种质创新国家重点实验室, 南京 210095
³荆楚理工学院, 特色花卉生物育种湖北省工程研究中心, 荆门 448000

收稿日期:2022-10-03 接受日期:2023-03-07 出版日期:2023-11-01 发布日期:2023-11-27
通讯作者: * E-mail: hyh2010@mail.hzau.edu.cn
作者简介:^† 共同第一作者
基金资助:
国家自然科学基金(32172616);特色花卉生物育种湖北省工程研究中心开放课题(2022ZD007)

Morphological Identification and Development of Linkage Markers for Lobed Ray Floret Mutants in Marigold (Tagetes erecta)

Wenjing Wang¹^,^†, Yu Zhu¹^,^†, Hongming Zhang¹^,², Ludan Wei¹, Qingping Yi³, Xiaomin Yu¹, Yuhan Liu¹, Lixue Zhang¹, Wenhan Cheng³, Yanhong He¹^,^*()

¹National Key Laboratory for Germplasm Innovation & Utilization of Horticultural Crops, College of Horticulture and Forestry Sciences, Huazhong Agricultural University, Wuhan 430070, China
²State Key Laboratory of Crop Genetics and Germplasm Enhancement, College of Horticulture, Nanjing Agricultural University, Nanjing 210095, China
³Hubei Engineering Research Center for Specialty Flowers Biological Breeding, Jingchu University of Technology, Jingmen 448000, China

Received:2022-10-03 Accepted:2023-03-07 Online:2023-11-01 Published:2023-11-27
Contact: * E-mail: hyh2010@mail.hzau.edu.cn
About author:^† These authors contributed equally to this paper

摘要/Abstract

摘要： 万寿菊(Tagetes erecta)是菊科万寿菊属重要的观赏花卉, 其舌状花花冠边缘的类型有平滑(smooth)、波浪(undulate)及不同程度的缺刻(incision)等。以第1轮小花花冠筒边缘五裂且雄蕊发育正常的万寿菊突变体JH和舌状花花冠边缘平滑且雄蕊败育的万寿菊自交系S5为亲本构建F₂代分离群体。遗传分析表明, JH突变性状由显性单基因控制, 将其命名为Telf。组织学和细胞学观察发现JH第1轮小花为舌状花管瓣化, 其花冠筒顶部具有5个裂片, 中下部融合呈管状, 且花药和花粉粒发育恢复正常。利用BSR-seq和比较基因组学方法, 将控制舌状花花冠裂片形成的基因Telf定位于37003-SCAR标记和34032-CAPS标记之间, 2个标记距Telf的遗传距离分别为3.684 cM和3.517 cM。该研究为后续精细定位目的基因Telf奠定了基础, 也为万寿菊分子标记开发提供了新方法。

关键词: 万寿菊, 舌状花, 裂片, BSR-seq, 分子标记

Abstract: Marigold (Tagetes erecta) is an important ornamental plant of the genus Tagetes in the Asteraceae. The morphology of its ray floret corolla varies from smooth, undulate to different degree of incision. An F₂ population was constructed from a cross between the mutant marigold JH with five-lobed corollas and fertile stamens in the first round florets and the inbred line S5 with smooth corolla of ray florets and aborting stamens. Inheritance analysis showed that the mutation trait of JH was controlled by a single-dominant gene, named Tagetes erecta lobed ray florets (Telf). Histological and cytological observations showed that the first round ray florets of JH were disk-like, with five lobes formed at the top of the corolla tube, the middle and lower parts were tubular, and the development of anthers and pollen grains are to normal. Using BSR-seq and comparative genomics methods, the gene Telf controlling lobed ray florets was mapped in the interval of 37003-SCAR and 34032-CAPS. 37003-SCAR marker was 3.684 cM from the Telf gene. On the other side of the gene, 34032-CAPS marker was placed about 3.517 cM from Telf. This study laid a foundation for the subsequent fine mapping of the target gene Telf, and also provided a new method for the selection of molecular markers in marigold.

Key words: Tagetes erecta, ray floret, lobed corolla, BSR-seq, molecular marker

王文静, 朱钰, 张洪铭, 韦陆丹, 易庆平, 余晓敏, 刘雨菡, 张莉雪, 程文翰, 何燕红. 万寿菊舌状花花冠裂片突变体的形态鉴定及连锁标记开发. 植物学报, 2023, 58(6): 893-904.

Wenjing Wang, Yu Zhu, Hongming Zhang, Ludan Wei, Qingping Yi, Xiaomin Yu, Yuhan Liu, Lixue Zhang, Wenhan Cheng, Yanhong He. Morphological Identification and Development of Linkage Markers for Lobed Ray Floret Mutants in Marigold (Tagetes erecta). Chinese Bulletin of Botany, 2023, 58(6): 893-904.

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链接本文: https://www.chinbullbotany.com/CN/10.11983/CBB22231

https://www.chinbullbotany.com/CN/Y2023/V58/I6/893

图/表 11

图1 万寿菊S5和JH的F2代群体构建及舌状花花冠裂片性状分离 Pi: 雌蕊; St: 雄蕊。Bars=1 cm

Figure 1 Construction of F2 population between marigold S5 and JH, and segregation of ray corolla traits in the F2 population Pi: Pistil; St: Stamen. Bars=1 cm

表1 酶切扩增多态性序列(CAPS)标记引物

Table 1 Primers for amplified polymorphic sequences (CAPS) markers

Unigene ID	Primer name	Forward primer (5'-3')	Reverse primer (5'-3')	Restriction enzyme
T01.PB44152	44152-CAPS	GTCAGCATGGGAATTCTACTGTTGG	CCATTTATAGCTGGAAAACAACCCG	DraI
T01.PB49370	49370-CAPS	TCGTCTCCACCATAACCACCGC	CTCAACTTTATCCGCGCCAATA	XhoI
T01.PB23497	23497-CAPS	TTCGTCCAAACCCAGATATGCTTAT	ACCAACCGACAAGTACGATTCGTAT	XbaI
T01.PB40376	40376-CAPS	GTGTGCGTCTCTGGCACCATCAT	TAAGTACCATTCTCCGACCCGCA	DraI
T01.PB44341	44341-CAPS	CCAACCTTTTTGTGCTTGCTAAC	AAGTTCCAAGTCTGTTACTTCCATC	XbaI
T01.PB36031	36031-CAPS	CGGTGACATTATGAACCTTGGACTG	CCCATAACAACAACTTAGTACCGTATT	EcoRI
T01.PB21624	21624-CAPS	CCGAGCATATTGAATTTTGGGTTAT	AGGGAATGAGAAGAACAATGGTCGT	SalI
T01.PB37003	37003-CAPS	TGACTCTTTCTATCGCAGAGCCACT	ATCCAGGTACAAGTCACATGTCTGT	EcoRI
T01.PB34032	34032-CAPS	GAGGTCTGCTTGCTTTTATGAGTAC	GCAAATATGGGCTACAAAATTCTATT	EcoRI
T01.PB26228	26228-CAPS	AATAGTCCAGCAGCAAAAGTTGGTC	GCTTTCAGTGAACCAAAGGATATGG	XbaI
T01.PB17555	17555-CAPS	TGCTCATACATCCATCTCCGTCTTC	CAGCTTTCTTGAACGTGTCCCTAAC	XbaI
T01.PB43155	43155-CAPS	GTATATGCTGCAACTCTGCCCAAAG	GGAACACGATCCATGTCAGGTG	EcoRI
T01.PB17261	17261-CAPS	GAGAAATACGAACCGTTATGG	AACTAGTACAACATTTTAAACCTGG	EcoRI
T01.PB40072	40072-CAPS	ATGCAGATGACCCTACAAATTACCT	GCCTCTATTCCTATTTTCTCCTACC	SalI

图2 万寿菊S5和JH盛花期花器官的体式显微镜和扫描电镜观察

Figure 2 Stereomicroscope and scanning electron microscope observation of floral organs of marigold S5 and JH at full blooming stage

图3 万寿菊S5和JH盛花期4轮花器官的半薄切片横切面观察 Vb: 维管束; Bu-like: 泡状细胞; UE: 上表皮细胞; ML: 中层组织; LE: 下表皮细胞; Po: 花粉粒; Pa: 乳突细胞。Bars=25 μm

Figure 3 The observation of transverse semi-thin sections in four whorls floral organs of marigold S5 and JH at full blooming stage Vb: Vascular bundles; Bu-like: Bubble-like cells; UE: Upper epidermis; ML: Middle layer; LE: Lower epidermis; Po: Pollen grains; Pa: Papilla cells. Bars=25 μm

图4 万寿菊S5 (A-E)和JH (F-J)不同发育时期花蕾的体式显微镜观察 (A), (F) 1 mm花蕾; (B), (G) 1.5 mm花蕾; (C), (H) 2 mm花蕾; (D), (I) 3 mm花蕾; (E), (J) 4 mm花蕾。IP: 花序原基; RFP: 舌状花原基; DFP: 管状花原基; Disc-like: 类似管状花花冠; Ray-like: 类似舌状花花冠。Bars=200 μm

Figure 4 Stereomicroscope observation of floral buds of marigold S5 (A-E) and JH (F-J) at different developmental stages (A), (F) 1 mm buds; (B), (G) 1.5 mm buds; (C), (H) 2 mm buds; (D), (I) 3 mm buds; (E), (J) 4 mm buds. IP: Inflorescence primordium; RFP: Ray floret primordium; DFP: Disc floret primordium; Disc-like: Disc-like floret corolla; Ray-like: Ray-like floret corolla. Bars=200 μm

图5 万寿菊S5 (A-F)和JH (G-L)不同发育时期花蕾石蜡切片横切面 (A), (G) 0.5 mm花蕾; (B), (H) 1 mm花蕾; (C), (I) 1.5 mm花蕾; (D), (J) 2 mm花蕾; (E), (K) 3 mm花蕾; (F), (L) 4 mm花蕾。IP: 花序原基; RFP: 舌状花原基; DFP: 管状花原基; PaP: 冠毛原基; CoRP: 舌状花花冠原基; CoDP: 管状花花冠原基; StP: 雄蕊原基; PiP: 雌蕊原基; Pa: 冠毛; Co: 花冠; St: 雄蕊; Pi: 雌蕊。Bars=100 μm

Figure 5 The transverse section of paraffin sections of floral buds of marigold S5 (A-F) and JH (G-L) at different developmental stages (A), (G) 0.5 mm buds; (B), (H) 1 mm buds; (C), (I) 1.5 mm buds; (D), (J) 2 mm buds; (E), (K) 3 mm buds; (F), (L) 4 mm buds. IP: Inflorescence primordium; RFP: Ray floret primordium; DFP: Disc floret primordium; PaP: Pappus primordium; CoRP: Corolla primordium of ray floret; CoDP: Corolla primordium of disc floret; StP: Stamen primordium; PiP: Pistil primordium; Pa: Pappus; Co: Corolla; St: Stamen; Pi: Pistil. Bars=100 μm

表2 BSR-seq质量

Table 2 Quality of BSR-seq

	S5-pool	JH-pool	F₂-S5	F₂-JH
Total reads	75274490	93514368	68385858	74073254
Clean reads	72943532	90821192	66109078	71708520
Total base	11291173500	14027155200	10257878700	11110988100
Clean base	10838757206	13499502354	9820452668	10648842550
GC content (%)	42	43	43	43
Q20 (%)	99.99	100	100	100
Q30 (%)	99.57	99.63	99.47	99.55

图6 基于BSR-seq分析结果的万寿菊转录组与向日葵基因组同源基因比对散点图灰点: 万寿菊基因在向日葵基因组中同源基因的位置; 彩色点: ΔSNP index≥0.3的万寿菊基因

Figure 6 Scatter diagram of homologous gene alignment between marigold transcriptome and sunflower genome based on the results of BSR-seq analysis Grey dots: Locations of homologous genes in marigold mapped to sunflower; Colorful dots: Unigenes in marigold with ΔSNP index≥0.3

表3 与万寿菊舌状花花冠裂片相关的SNP位点信息

Table 3 The SNPs associated with lobes of ray florets in marigold

Unigene ID	Primer name	Position	ΔSNP index	Mutant position	Mutant base
T01.PB44152	44152-CAPS	32539231	0.873	2113	A→T
T01.PB49370	49370-CAPS	50400088	0.763	144	G→T
T01.PB23497	23497-CAPS	138318439	0.555	975	A→G
T01.PB40376	40376-CAPS	158179471	0.751	402	T→A
T01.PB44341	44341-CAPS	174818902	0.666	225	T→C
T01.PB36031	36031-CAPS	176617357	0.715	1712	A→G
T01.PB21624	21624-CAPS	189237745	0.630	220	G→C
T01.PB37003	37003-SCAR	191264434	0.711	1054	T→A
T01.PB34032	34032-CAPS	199916490	0.846	1138	G→A
T01.PB26228	26228-CAPS	204195845	0.533	1471	C→T
T01.PB17555	17555-CAPS	208494177	0.740	343	C→T
T01.PB43155	43155-CAPS	210482932	0.653	1475	T→C
T01.PB17261	17261-CAPS	238981210	0.553	1316	A→C
T01.PB40072	40072-CAPS	250623369	0.555	2109	C→T

表4 万寿菊8个基因的分子标记交换单株分布

Table 4 The individuals with the exchange of molecular markers for eight genes in marigold

CAPS markers	F₂-S5
CAPS markers	1	2	3	4	5	6	7	8	9	10	11	12	13	14	15	16	17	18	19	20	21	22	23	24	25	26	27	28	29	30
36031-CAPS					*										*									*				*
21624-CAPS																	*
37003-SCAR																	*
34032-CAPS																					*									*
44152-CAPS		*															*				*
49370-CAPS		*															*				*
17555-CAPS		*															*	*			*
43155-CAPS		*															*	*			*
CAPS markers	F₂-JH
CAPS markers	1	2	3	4	5	6	7	8	9	10	11	12	13	14	15	16	17	18	19	20	21	22	23	24	25	26	27	28	29	30
36031-CAPS		*					*
21624-CAPS										*
37003-SCAR
34032-CAPS		*
44152-CAPS										*
49370-CAPS										*
17555-CAPS										*														*
43155-CAPS										*														*

图7 37003-SCAR标记(A)和34032-CAPS标记(B)在亲本和F2代群体中的PCR扩增结果 M: 2 000 bp marker; 1-10: 舌状花花冠无裂片以及无雄蕊的F2代单株; 11-20: 舌状花花冠有5个裂片以及有雄蕊的F2代单株, 其中11-15为纯合子, 16-20为杂合子。

Figure 7 PCR amplification results of 37003-SCAR marker (A) and 34032-CAPS marker (B) in parents and F2 population M: 2 000 bp marker; 1-10: Individuals of F2 with ray floret without lobes and stamens; 11-20: Individuals of F2 that have ray florets with five-lobed and stamens, of which 11-15 are homozygotes and 16-20 are heterozygotes.

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