白色桑椹颜色缺失分子机制初步探讨

doi:10.16420/j.issn.0513-353x.2024-0929

摘要/Abstract

摘要：

为揭示白色桑椹中花青素苷合成阻断的机理，对白色桑椹和黑色桑椹的代谢组与转录组进行了比较和分析。结果表明，‘安椹’（黑色果实）在成熟过程中花青素苷含量逐步积累，在成熟期积累量最大；‘桂花’（白色果实）在青果期花青素苷含量最高，在转色期和成熟期的花青素苷含量较低。花青素苷关键呈色物质矢车菊素、飞燕草素、天竺葵素、锦葵色素、矮牵牛素等均在‘桂花’中积累量较低，相反矢车菊素-3-O-葡萄糖苷、矢车菊素-3-O-芸香糖苷、飞燕草素-3-O-葡萄糖苷、飞燕草素-3-O-槐糖苷、天竺葵素-3-O-葡萄糖苷、天竺葵素-3-O-芸香糖苷、原花青素B1、原花青素B3等物质在‘安椹’中大量积累。转色期‘桂花’中参与花青素苷生物合成的PAL、C4H、4CL、CHS、CHI、F3H等基因表达减少，这种下调阻止了花青素苷在白色桑椹中的积累。基于转录组数据鉴定到6个差异表达的MYB转录因子基因，结合基因表达、进化分析以及下游靶基因鉴定，分析MYB6可能在白色桑椹成熟过程中花青素苷的合成中起抑制作用。

关键词: 桑椹, 转录组, 代谢组, 花青素, 结构基因, MYB6

Abstract:

In order to reveal the mechanism of anthocyanin synthesis blockade in white mulberry，the metabolome and transcriptome of white mulberry and black mulberry were compared and analyzed. The results showed that the anthocyanin content of‘Anshen’（black fruit）gradually accumulated during the ripening process，and the accumulation amount was the highest in the ripening stage，while the anthocyanin content of‘Guihua’（white fruit）was the highest in the green fruit stage，and the anthocyanin content was lower at the veraison and maturity stages. The key color substances of anthocyanin glycosides，such as cyanidin，delphinin，pelargondin，malvidin pigment and petunidin，all had low accumulation in‘Guihua’，especially cyanidin-3-O-glucoside，cyanidin-3-O-rutinoside，delphin-3-O-glucoside，delphin-3-O-loccoside，pelargondin-3-O-glucoside，pelargondin-3-O-rutinoside，proanthocyanidin B1，proanthocyanidinin B3 and other substances showed a large accumulation in‘Anshen’. The expression of PAL，C4H，4CL，CHS，CHI，F3H and other genes involved in the biosynthesis of anthocyanins in‘Guihua’decreased during veraison，and this down-regulation prevented the accumulation of anthocyanins in white mulberry. Based on transcriptome data，six differentially expressed MYB transcription factor genes were identified，and combined with gene expression，evolutionary analysis and downstream target gene identification，it was analyzed that MYB6 may play an inhibitory role in the synthesis of anthocyanin glycosides during the maturation of white mulberry.

Key words: mulberry, transcriptome, metabolome, anthocyanin, structural gene, MYB6

郭新淼, 田露瑶, 曹佳琪, 谢岩, 高妍夏, 孙志超. 白色桑椹颜色缺失分子机制初步探讨[J]. 园艺学报, 2025, 52(6): 1451-1462.

GUO Xinmiao, TIAN Luyao, CAO Jiaqi, XIE Yan, GAO Yanxia, and SUN Zhichao. Preliminary Research on the Molecular Mechanism of Color Loss in White Mulberry Fruit[J]. Acta Horticulturae Sinica, 2025, 52(6): 1451-1462.

图/表 6

参考文献 33

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物质 Compound	桂花S2 Guihua S2	安椹S2 Anshen S2
矢车菊素-3-O-对香豆酰葡萄糖苷Cyanidin-3-O-(coumaryl)-glucoside	0.66 ± 0.04 a	0.98 ± 0.19 a
矢车菊素-3-O-葡萄糖苷Cyanidin-3-O-glucoside	2.68 ± 1.51 B	192.33 ± 32.01 A
矢车菊素-3-O-芸香糖苷Cyanidin-3-O-rutinoside	1.05 ± 0.37 B	188.33 ± 9.61 A
矢车菊素-3-O-槐糖苷 Cyanidin-3-O-sophoroside	0.03 ± 0.02 b	0.58 ± 0.10 a
矢车菊素-3-O-丙二酰葡萄糖苷Cyanidin-3-O-(6-O-malonyl-beta-D-glucoside)	0	0.01 ± 0
矢车菊素-3,5,3'-O-三葡萄糖苷 Cyanidin-3,5,3′-O-triglucoside	0.01 ± 0	0
飞燕草素-3-O-葡萄糖苷Delphinidin-3-O-glucoside	2.12 ± 0.14 B	7.37 ± 0.35 A
飞燕草素-3-O-丙二酰葡萄糖苷Delphinidin-3-O-(6-O-malonyl-beta-D-glucoside)	0.23 ± 0.01 B	1.33 ± 0.11 A
飞燕草素-3-O-槐糖苷 Delphinidin-3-O-sophoroside	1.68 ± 0.25 B	10.95 ± 1.73 A
飞燕草素-3-芸香糖苷-5-葡萄糖苷Delphinidin-3-rutinoside-5-glucoside	0.42 ± 0.04 B	0.86 ± 0.07 A
飞燕草素-3-O-半乳糖苷 Delphinidin-3-O-galactoside	0.05 ± 0.01 a	0.02 ± 0 b
飞燕草素-3-O-(6-O-丙二酰基)-葡萄糖苷-3′-葡萄糖基 Delphinidin-3-O-(6-O-malonyl)-glucoside-3′-glucoside	0.02 ± 0 B	0.06 ± 0.01 A
天竺葵素-3-O-葡萄糖苷Pelargonidin-3-O-glucoside	0.26 ± 0.14 B	4.81 ± 0.82 A
天竺葵素-3-O-芸香糖苷 Pelargonidin-3-O-rutinoside	0.05 ± 0.02 B	4.02 ± 0.41 A
天竺葵素-3-O-槐糖苷 Pelargonidin-3-O-sophoroside	0.02 ± 0	0.03 ± 0.01
天竺葵素-3-O-丙二酰葡萄糖苷Pelargonidin-3-O-(6-O-malonyl-beta-D-glucoside)	0.01 ± 0 b	0.01 ± 0 a
矮牵牛素-3-O-芸香糖苷 Petunidin-3-O-rutinoside	0	0
矮牵牛素-3-O-丙二酰葡萄糖苷Petunidin-3-O-(6-O-malonyl-beta-D-glucoside)	0	0
矮牵牛素-3-O-葡萄糖Petunidin-3-O-glucoside	0.01 ± 0 b	0.02 ± 0 a
锦葵色素-3-O-葡萄糖苷Malvidin-3-O-glucoside	0.03 ± 0 B	0.06 ± 0.01 A
芍药花色素-3-O-葡萄糖苷 Peonidin-3-O-glucoside	0	0.10 ± 0.01
芍药花色素-3-O-芸香糖苷 Peonidin-3-O-rutinoside	0.01 ± 0	0.01 ± 0
原花青素A1 Procyanidin A1	0.05 ± 0.20	0
原花青素B1 Procyanidin B1	0.04 ± 0.01 B	19.60 ± 1.87 A
原花青素B2 Procyanidin B2	0	0.07 ± 0.03
原花青素B3 Procyanidin B3	0.10 ± 0.04 B	51.33 ± 3.87 A
柚皮素-7-O-葡萄糖苷 Naringenin-7-O-glucoside	0	0.11 ± 0.018
柚皮素 Naringenin	0.03 ± 0.01 B	0.06 ± 0 A
芦丁 Rutin	363.00 ± 22.72 B	1270.00 ± 55.68 A
香橙素（二氢山奈酚）Dihydrokaempferol	0.16 ± 0.03 a	0.18 ± 0.01 a
山柰酚-3-O-芸香糖苷Kaempferol-3-O-rutinoside	42.50 ± 3.759 a	76.43 ± 15.37 a
异槲皮苷Quercetin-3-O-glucoside	143.00 ± 8.19 B	485.67 ± 21.08 A

物质 Compound	桂花S2 Guihua S2	安椹S2 Anshen S2
矢车菊素-3-O-对香豆酰葡萄糖苷Cyanidin-3-O-(coumaryl)-glucoside	0.66 ± 0.04 a	0.98 ± 0.19 a
矢车菊素-3-O-葡萄糖苷Cyanidin-3-O-glucoside	2.68 ± 1.51 B	192.33 ± 32.01 A
矢车菊素-3-O-芸香糖苷Cyanidin-3-O-rutinoside	1.05 ± 0.37 B	188.33 ± 9.61 A
矢车菊素-3-O-槐糖苷 Cyanidin-3-O-sophoroside	0.03 ± 0.02 b	0.58 ± 0.10 a
矢车菊素-3-O-丙二酰葡萄糖苷Cyanidin-3-O-(6-O-malonyl-beta-D-glucoside)	0	0.01 ± 0
矢车菊素-3,5,3'-O-三葡萄糖苷 Cyanidin-3,5,3′-O-triglucoside	0.01 ± 0	0
飞燕草素-3-O-葡萄糖苷Delphinidin-3-O-glucoside	2.12 ± 0.14 B	7.37 ± 0.35 A
飞燕草素-3-O-丙二酰葡萄糖苷Delphinidin-3-O-(6-O-malonyl-beta-D-glucoside)	0.23 ± 0.01 B	1.33 ± 0.11 A
飞燕草素-3-O-槐糖苷 Delphinidin-3-O-sophoroside	1.68 ± 0.25 B	10.95 ± 1.73 A
飞燕草素-3-芸香糖苷-5-葡萄糖苷Delphinidin-3-rutinoside-5-glucoside	0.42 ± 0.04 B	0.86 ± 0.07 A
飞燕草素-3-O-半乳糖苷 Delphinidin-3-O-galactoside	0.05 ± 0.01 a	0.02 ± 0 b
飞燕草素-3-O-(6-O-丙二酰基)-葡萄糖苷-3′-葡萄糖基 Delphinidin-3-O-(6-O-malonyl)-glucoside-3′-glucoside	0.02 ± 0 B	0.06 ± 0.01 A
天竺葵素-3-O-葡萄糖苷Pelargonidin-3-O-glucoside	0.26 ± 0.14 B	4.81 ± 0.82 A
天竺葵素-3-O-芸香糖苷 Pelargonidin-3-O-rutinoside	0.05 ± 0.02 B	4.02 ± 0.41 A
天竺葵素-3-O-槐糖苷 Pelargonidin-3-O-sophoroside	0.02 ± 0	0.03 ± 0.01
天竺葵素-3-O-丙二酰葡萄糖苷Pelargonidin-3-O-(6-O-malonyl-beta-D-glucoside)	0.01 ± 0 b	0.01 ± 0 a
矮牵牛素-3-O-芸香糖苷 Petunidin-3-O-rutinoside	0	0
矮牵牛素-3-O-丙二酰葡萄糖苷Petunidin-3-O-(6-O-malonyl-beta-D-glucoside)	0	0
矮牵牛素-3-O-葡萄糖Petunidin-3-O-glucoside	0.01 ± 0 b	0.02 ± 0 a
锦葵色素-3-O-葡萄糖苷Malvidin-3-O-glucoside	0.03 ± 0 B	0.06 ± 0.01 A
芍药花色素-3-O-葡萄糖苷 Peonidin-3-O-glucoside	0	0.10 ± 0.01
芍药花色素-3-O-芸香糖苷 Peonidin-3-O-rutinoside	0.01 ± 0	0.01 ± 0
原花青素A1 Procyanidin A1	0.05 ± 0.20	0
原花青素B1 Procyanidin B1	0.04 ± 0.01 B	19.60 ± 1.87 A
原花青素B2 Procyanidin B2	0	0.07 ± 0.03
原花青素B3 Procyanidin B3	0.10 ± 0.04 B	51.33 ± 3.87 A
柚皮素-7-O-葡萄糖苷 Naringenin-7-O-glucoside	0	0.11 ± 0.018
柚皮素 Naringenin	0.03 ± 0.01 B	0.06 ± 0 A
芦丁 Rutin	363.00 ± 22.72 B	1270.00 ± 55.68 A
香橙素（二氢山奈酚）Dihydrokaempferol	0.16 ± 0.03 a	0.18 ± 0.01 a
山柰酚-3-O-芸香糖苷Kaempferol-3-O-rutinoside	42.50 ± 3.759 a	76.43 ± 15.37 a
异槲皮苷Quercetin-3-O-glucoside	143.00 ± 8.19 B	485.67 ± 21.08 A

转录因子结合位点 Transcription factor binding site	靶向序列名字 Sequence name	链 Strand	起始位置/bp Start site	结束位置/bp End site	P值 P-value	Q值 Q -value	匹配序列 Matched sequence
GCTTGTTG	OMT（LOC21410121）	+	1 265	1 272	1.47E-05	0.383	GCTTGTTG
GCTTGTTG	OMT（LOC21410121）	+	1 568	1 575	1.47E-05	0.383	GCTTGTTG
rymAGTTA	PAL（LOC21407112）	+	1 922	1 929	6.83E-05	1	GCAAGTTA
dwmwGTTr	OMT（LOC21410121）	-	673	680	5.87E-05	1	GTCAGTTG
dwmwGTTr	UFGT（LOC21403843）	-	1 386	1 393	5.87E-05	1	GACTGTTG
wwmaGTTr	PAL（LOC21407114）	+	921	928	7.15E-05	1	TTCAGTTG
rsTwGkTr	C4H（LOC213945240）	+	660	667	4.82E-05	0.417	GGTAGGTG
	CHI（LOC21400267）	-	1 491	1 498	4.82E-05	0.417	GCTAGGTG
	4CL（LOC21402620）	+	1 507	1 514	4.82E-05	0.417	GGTAGGTG
	4CL（LOC21402620）	-	1 744	1 751	4.82E-05	0.417	GGTAGGTG
	C4H（LOC213945240）	-	1 825	1 832	4.82E-05	0.417	GGTTGGTG
	PAL（LOC21407114）	-	1 967	1 974	4.82E-05	0.417	GGTTGGTG
rbTrGTTA	CHS（LOC21400913）	-	1 740	1 747	2.94E-05	1	GCTGGTTA

转录因子结合位点 Transcription factor binding site	靶向序列名字 Sequence name	链 Strand	起始位置/bp Start site	结束位置/bp End site	P值 P-value	Q值 Q -value	匹配序列 Matched sequence
GCTTGTTG	OMT（LOC21410121）	+	1 265	1 272	1.47E-05	0.383	GCTTGTTG
GCTTGTTG	OMT（LOC21410121）	+	1 568	1 575	1.47E-05	0.383	GCTTGTTG
rymAGTTA	PAL（LOC21407112）	+	1 922	1 929	6.83E-05	1	GCAAGTTA
dwmwGTTr	OMT（LOC21410121）	-	673	680	5.87E-05	1	GTCAGTTG
dwmwGTTr	UFGT（LOC21403843）	-	1 386	1 393	5.87E-05	1	GACTGTTG
wwmaGTTr	PAL（LOC21407114）	+	921	928	7.15E-05	1	TTCAGTTG
rsTwGkTr	C4H（LOC213945240）	+	660	667	4.82E-05	0.417	GGTAGGTG
	CHI（LOC21400267）	-	1 491	1 498	4.82E-05	0.417	GCTAGGTG
	4CL（LOC21402620）	+	1 507	1 514	4.82E-05	0.417	GGTAGGTG
	4CL（LOC21402620）	-	1 744	1 751	4.82E-05	0.417	GGTAGGTG
	C4H（LOC213945240）	-	1 825	1 832	4.82E-05	0.417	GGTTGGTG
	PAL（LOC21407114）	-	1 967	1 974	4.82E-05	0.417	GGTTGGTG
rbTrGTTA	CHS（LOC21400913）	-	1 740	1 747	2.94E-05	1	GCTGGTTA