Scientia Agricultura Sinica ›› 2023, Vol. 56 ›› Issue (3): 416-429.doi: 10.3864/j.issn.0578-1752.2023.03.002

• CROP GENETICS & BREEDING·GERMPLASM RESOURCES·MOLECULAR GENETICS • Previous Articles     Next Articles

Cloning of MYBL2 Gene from Brassica and Its PCR Identification in Genomes A, B and C

ZOU Ting1(), LIU LiLi1, XIANG JianHua1, ZHOU DingGang1, WU JinFeng1, LI Mei2, LI Bao2, ZHANG DaWei1(), YAN MingLi1,2()   

  1. 1College of Life Science and Health, Hunan University of Science and Technology/Hunan Provincial Key Laboratory of Genetic Improvement and Comprehensive Utilization of Cash Crops, Xiangtan 410201, Hunan
    2Hunan Crop Research Institute, Changsha 410125
  • Received:2022-08-31 Accepted:2022-10-24 Online:2023-02-01 Published:2023-02-14
  • Contact: ZHANG DaWei, YAN MingLi E-mail:3392648002@qq.com;zhangdawei.hnust@foxmail.com;ymljack@126.com

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Abstract:

【Objective】In Arabidopsis, MYBL2 negatively regulates the biosynthesis of anthocyanins and proanthocyanidins. The MYBL2 genes from six Brassica species with different leaf colors were cloned. By analyzing the sequence and expression pattern of MYBL2, the function of MYBL2 in the biosynthesis of anthocyanins in Brassica species was explored. 【Method】The sequences of MYBL2 from the six Brassica species with different leaf colors were obtained using homology-based cloning method and multi sequence alignment and phylogenetic tree analysis were performed. The purple leaf materials of B. rapa, B. napus, B. juncea and B. carinata were treated with shading, and the expression level of MYBL2 gene was analyzed by transcriptome sequencing and qRT-PCR. The qRT-PCR in B. oleracea, B. napus, B. juncea and B. carinata with purple and green leaves were also performed to evaluate the expression level of MYBL2. Based on the nucleotide variation sites of the cloned MYBL2-1 and MYBL2-2 sequences, PCR markers which could distinguish the genomic origin of alleles were developed. 【Result】A total of 56 copies of 9 homologs of MYBL2-1 and MYBL2-2 were cloned from 19 samples of six species of Brassica. The BcaMYBL2-1 gene was obtained for the first time. The total length of BcaMYBL2-1 sequence was 867 bp, including two introns of 168 bp and 102 bp respectively, encoding 198 amino acids, with a molecular weight of 22.69 kD and an isoelectric point (pI) of 8.72. Sequence alignment and evolutionary analysis showed that BcaMYBL2-1 was derived from B genome. Among the MYBL2-1 and MYBL2-2 copies of six species in Brassica, only BraA07.MYBL2-1, BolC06.MYBL2-1 and BcaMYBL2-1 exhibited sequence differences in different leaf color materials. After shading treatment, the leaf color of purple leaf material becomes lighter than that of the unshaded part. In Chinese cabbage Zibao 5, the expression of BraA07.MYBL2-1 and BraA02.MYBL2-2 in the shaded part were 0.7 and 0.4 times of that in the unshaded part, respectively. In Brassica napus with purple leaves and white flowers, the expression of BnaA07.MYBL2-1, BnaC06.MYBL2-1, BnaA02.MYBL2-2 and BnaC02.MYBL2-2 in the shaded part were 0.4, 0.5, 0.4 and 0.4 times of that in the unshared part, respectively. In purple leaf mustard, the expression of BjuA07.MYBL2-1, BjuB03.MYBL2-1, BjuA02.MYBL2-2 and BjuB05.MYBL2-2 in the shaded part were 0.4, 0.3, 0.4 and 0.2 times of that in the unshared part, respectively. In B. carinata with purple leaf, the expression of BcaMYBL2-1, BcaB03.MYBL2-1 and BcaC03.MYBL2-2 in the shaded part were 0.3, 0.4 and 0.5 times of that in the unshared part, respectively. However, the expression of BcaB05.MYBL2-2 in the shaded part was 2.4 times of that in the unshared part. Comparing the expression of MYBL2 gene in different leaf color materials of Brassica, the results showed that the expression of most of the homologous genes of MYBL2 gene in purple leaf materials was higher than that in green leaf materials except kale. In kale, the expression of BolC06.MYBL2-1 and BolC02.MYBL2-2 in green kale were 2.5 and 3.5 times that in purple kale, respectively. In Brassica napus, the expression of BnaA07.MYBL2-1, BnaC06.MYBL2-1 and BnaC02.MYBL2-2 in purple leaf and white flower were 7.5, 8.6 and 26.0 times of that in green leaf and white flower, while the expression of BnaA02.MYBL2-2 in green leaf and white flower was 13.0 times of that in purple leaf and white flower. The expression levels of BjuA07.MYBL2-1, BjuB03.MYBL2-1, BjuA02.MYBL2-2 and BjuB05.MYBL2-2 in Brassica juncea were 8.3 times, 11.8 times, 23.2 times and 14.6 times of those in Sichuan yellow respectively. In B. carinata with purple leaf, BcaMYBL2-1 and BcaB03.MYBL2-1 were 7.1 and 27.6 times as much as W-BCDH76, respectively. However, BcaB05.MYBL2-2 and BcaC03.MYBL2-2 genes of W-BCDH76 were 2.8 and 5.0 times as much as those of B. carinata with purple leaf, respectively. Five pairs of primers were obtained, which can effectively identify the MYBL2 from A, B and C genomes of Brassica. 【Conclusion】After shading treatment, MYBL2 gene expression was closely related to light. The regulation mechanism of MYBL2 gene in Brassica plants involved in anthocyanin biosynthesis was different from that of Arabidopsis plants in which MYBL2 gene negatively regulated anthocyanin biosynthesis.

Key words: Brassica species, MYBL2 gene, homologous cloning, gene expression, genomic PCR identification

Fig. 1

Amino acid sequence alignment of genes encoding BniB03.MYBL2-1, BcaMYBL2-1 and BjuB03.MYBL2-1 of different leaf color materials BniB03 G: B.nigra (Giebra); Bca G:WBCDH76; Bca P: B. carinata with purple leaf; BjuB03 SY: Sichuan Yellow; BjuB03 PM: Purple leaf mustard"

Fig. 2

Phylogenetic tree constructed by different leaf color materials of 6 species of Brassica and amino acid sequences encoded by MYBL2 gene of other plants Bra G: B. rapa with green leaf; Bra P1: B. rapa with whole purple plant; Bra P2: Zibao 5; Bni G: Giebra; Bol G: Green leaf kale; Bol P: Purple leaf kale; SY: Sichuan Yellow; PM: Purple leaf mustard; Bna G: B. napus with green leaves and white flowers; Bna P: B. napus with purple leaves and white flowers; Bca G1: W-BCDH76; Bca G2: Huangai; Bca P: B. carinata with purple leaf; At: Arabidopsis thaliana; Rs: Raphanus sativus"

Table 1

Sequence difference of MYBL2 gene in Brassica with different leaf color materials"

材料名称
Material name		 类型
Type		 叶片颜色
Leaf color		 基因
Gene		 基因编码区核苷酸变异位点（ATG至TGA）
Sites of nucleotide polymorphism (bp)
57 495 540-600 646 751 842
绿叶白菜
B. rapa with green leaf		 白菜
B. rapa		 绿色
Green		 BraA07.MYBL2-1
 C G
 ...
 A
 T
 T
白菜全紫株
B. rapa with whole purple plant		 白菜
B. rapa		 紫色
Purple		 BraA07.MYBL2-1
 A T
 61
 G
 A
 A
紫宝5号
Zibao 5		 白菜
B. rapa		 紫色
Purple		 BraA07.MYBL2-1
 C G
 ...
 A T T
四川黄籽
Sichuan Yellow		 芥菜型油菜
B. juncea		 绿色
Green		 BjuA07.MYBL2-1
 A T
 61
 G
 A
 A
紫叶芥
Purple leaf mustard		 芥菜型油菜
B. juncea		 紫色
Purple		 BjuA07.MYBL2-1
 A T
 61
 G
 A
 A
绿叶白花
B. napus with green leaves and white flowers		 甘蓝型油菜
B. napus		 绿色
Green		 BnaA07.MYBL2-1
 A G
 ...
 A
 T
 T
紫叶白花
B. napus with purple leaves and white flowers		 甘蓝型油菜
B. napus		 紫色
Purple		 BnaA07.MYBL2-1
 A G
 ...
 A
 T
 T
4 18 67 217 268 395 415 519 684
W-BCDH76 埃塞俄比亚芥
B. carinata		 绿色
Green		 BcaMYBL2-1
 C
 C
 G
 T
 A
 C
 A
 G
 T
紫秆埃芥
B. carinata with purple leaf		 埃塞俄比亚芥
B. carinata		 紫色
Purple		 BcaMYBL2-1
 A
 T
 A
 C
 G
 T
 G
 T
 C
27 58 63 268 271 399-401 732 803 884
绿叶羽衣甘蓝
Green leaf kale		 甘蓝
B. oleracea		 绿色
Green		 BolC06.MYBL2-1
 A
 A
 ...
 C
 G
 TTG
 A
 G
 T
紫叶羽衣甘蓝
Purple leaf kale		 甘蓝
B. oleracea		 紫色
Purple		 BolC06.MYBL2-1
 C
 G
 T
 A
 A
 ...
 G
 T
 C

Fig. 3

The expression MYBL2 in Brassica with purple leaf after shading treatment A: Phenotype of Brassica purple leaf materials after shading treatment; B: Zibao 5; C: B. napus with purple leaf and white; D: Purple leaf mustard; E: B. carinata with purple leaf. sP: Untreated part of purple leaf; sG: Purple leaf shading treatment part"

Fig. 4

Heatmaps of differentially expressed genes (DEGs) expression pattern of anthocyanin biosynthesis after shading treatment of Brassica purple leaf materials A: Zibao 5; B: B. napus with purple leaves and white flowers; C: Purple leaf mustard; D: B. carinata with purple leaf; sP: Untreated part of purple leaf; sG: Purple leaf shading treatment part"

Fig. 5

The expression of MYBL2 in Brassica with different leaf color A: Phenotypes of Brassica with different leaf colors; B(P): Green leaf kale; B(G): Purple leaf kale; C(P): B. napus with purple leaves and white flowers; C(G): B. napus with green leaves and white flowers; D(P): Purple leaf mustard; D(G): Sichuan Yellow; E(P): B. carinata with purple leaf; E(G): W-BCDH76; P: Materials with purple leaf; G: Materials with green leaf"

Fig. 6

PCR detection of MYBL2-1 gene and MYBL2-2 gene from Brassica genomes A, B and C M: Marker; A: Zaoshu 40; B: B. nigra (Giebra); C: Kale; AB1: Purple leaf mustard; AB2: Li G19; AC1:Xiangyou787; AC2: B. napus with purple leaves and white flowers; BC1: Huangai; BC2: B. carinata with purple leaf; N: Negative control; ACT: Internal reference control"

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