草珊瑚中类胡萝卜素合成的内源激素调控机制分析

doi:10.13560/j.cnki.biotech.bull.1985.2023-1047

摘要/Abstract

摘要：

【目的】为了探究草珊瑚[Sarcandra glabra（thunb）nakai]不同器官中内源激素对类胡萝卜素差异积累的可能调控机制。【方法】通过代谢组学方法分析草珊瑚中内源激素和类胡萝卜素代谢物在不同器官间的差异分布情况，结合转录组学技术挖掘草珊瑚类胡萝卜素相关差异酶基因，进一步预测参与调控差异酶基因的转录因子及其激素相关顺式响应元件，从而分析内源激素对类胡萝卜素差异积累的可能调控作用。【结果】吲哚-3-羧酸（indole-3-carboxylic acid）、反式茉莉酸[（-）-jasmonic acid]、二氢茉莉酮酸甲酯（ethyl dihydrojasmonate）、油菜素甾酮（castasterone）、油菜素内酯（brassinolide）等7种内源激素在叶中的含量更高，与角黄素（canthaxanthin）、海胆酮（echinenone）、新黄质（neoxanthin）和念珠藻黄素（nostoxanthin）等8种差异代谢物等富集部位一致；而脱落酸（abscisic acid，ABA）、赤霉素4, 5-甲氧基水杨酸（gibberellin 4,5-methoxysalicylic acid）、二氢茉莉酸（dihydrojasmonic acid）和5,6-二羟基吲哚（5,6-dihydroxyindole）在根中的含量更高，与脱落酸醛（abscisic aldehyde）、4,4'-二聚戊烯（4,4'-aiapolycopenedial）、花药黄质（antheraxanthin）这3种差异代谢物富集部位一致；关键转录因子MYB106、SPL1、NAC015、ERF064、WRKY44、BHLH116可通过响应AUX、SA、GA、ABA、Me_JA等植物激素的刺激，参与调控类胡萝卜素合成途径的DXS、VDE、ABA2、AOG、ZEP、CYP97C1、DWARF27、CRTISO、LCYB、PSY这10种代谢酶的表达。实时荧光定量 PCR（quantitative real-time polymerase chain reaction, RT-qPCR）结果表明，随机选取的8个差异基因表达趋势与转录组测序结果一致。【结论】筛选出草珊瑚叶和根与类胡萝卜素相关的15 种差异内源激素与 11 种差异代谢物，预测了6种转录因子参与调控10 种代谢酶。

关键词: 草珊瑚, 类胡萝卜素, 转录组, 代谢组

Abstract:

【Objective】This work aims to explore the possible regulatory mechanism of endogenous hormones on the differential accumulation of carotenoids in different organs of Sarcandra glabra. 【Method】In this study, metabolomics was used to analyze the differential distribution of endogenous hormones and carotenoid metabolites in different organs of S. glabra. Combined with transcriptomics technology, the differential enzyme genes related to carotenoids in S. glabra were excavated, and the transcription factors involved in the regulation of differential enzyme genes and their hormone-related cis-response elements were further predicted, so as to analyze the possible regulation of endogenous hormones on the differential accumulation of carotenoids. 【Result】The contents of seven endogenous hormones such as indole-3-carboxylic acid,[（-）-jasmonic acid], ethyl dihydrojasmonate, castasterone and brassinolide were high in the leaves. Consistent with the enrichment sites of 8 differential metabolites, such as canthaxanthin, echinenone, neoxanthin and nostoxanthin; and abscisic acid, gibberellin 4,5-methoxysalicylic acid, dihydrojasmonic acid and 5,6-dihydroxyindole was high in the roots, it was consistent with the accumulation site of 3 differential metabolites, abscisic aldehyde, 4,4'-aiapolycopenedial and antheraxanthin. The key transcription factor MYB106, SPL1, NAC015, ERF064, WRKY44 and BHLH116 responded to the stimulation of plant hormones such as AUX, SA, GA, ABA, and Me_ JA, and participate in the regulation of carotenoid synthesis pathway DXS, VDE, ABA2, AOG, ZEP, CYP97C1, DWARF27, CRTISO, LCYB expression of these 9 metabolic enzymes. Quantitative real-time polymerase chain reaction（RT-qPCR）results showed that the expression trends of 8 randomly selected differentially expressed genes were consistent with the transcriptome sequencing results. 【Conclusion】The 15 differential endogenous hormones and 11 differential metabolites related to carotenoids in the leaves and roots of Sarcandra glabra were screened, and 6 transcription factors were predicted to be involved in the regulation of 16 metabolic enzymes.

Key words: Sarcandra glabra, carotenoids, transcriptome, metabolome

吴迪, 游小凤, 郑亦铮, 林楠, 张燕燕, 魏艺聪. 草珊瑚中类胡萝卜素合成的内源激素调控机制分析[J]. 生物技术通报, 2024, 40(5): 203-214.

WU Di, YOU Xiao-feng, ZHENG Yi-zheng, LIN Nan, ZHANG Yan-yan, WEI Yi-cong. Analysis of Endogenous Hormone Regulation Mechanism for Carotenoid Synthesis in Sarcandra glabra[J]. Biotechnology Bulletin, 2024, 40(5): 203-214.

图/表 11

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基因 Gene	正向引物 Forward primer（5'-3'）	反向引物Reverse primer（5'-3'）
CAC	TCCGACAAATTGGAGGTTGC	TGCTGCTGACAACAATCACG
DXS	GCCTTGAACGGACTCTTAGAC	AGGTGGAGAGCATCCTTGTT
PSY	GGCGTCCTTGTTGTTGTACC	ATGAGTGCTCTAGTGTAGGCTATG
NCED	CTCCAAGGCGTGTTCTATCG	CTCCAAGGCGTGTTCTATCG
VDE	CACGACAACGAGTACCTTCAC	TGACAGCACCACCATATCCA
PDS	ACGAGTACCAGGTGTTGTTCTA	AATCAGTTTGCCAGCATTCATTG
LYCB	GGAAGATATACAGGAGAGGATGGT	TCTGAGGAAGCACTGGAAGT
ZEP	GCTTGGAGGCAGAGTATTGAC	GCAGCCATTCTAGCCATTCC
CYP97C1	GATGAGAGCACAACAAGAGGTT	CGCCTTATTAACACTGGTGGAT

基因 Gene	正向引物 Forward primer（5'-3'）	反向引物Reverse primer（5'-3'）
CAC	TCCGACAAATTGGAGGTTGC	TGCTGCTGACAACAATCACG
DXS	GCCTTGAACGGACTCTTAGAC	AGGTGGAGAGCATCCTTGTT
PSY	GGCGTCCTTGTTGTTGTACC	ATGAGTGCTCTAGTGTAGGCTATG
NCED	CTCCAAGGCGTGTTCTATCG	CTCCAAGGCGTGTTCTATCG
VDE	CACGACAACGAGTACCTTCAC	TGACAGCACCACCATATCCA
PDS	ACGAGTACCAGGTGTTGTTCTA	AATCAGTTTGCCAGCATTCATTG
LYCB	GGAAGATATACAGGAGAGGATGGT	TCTGAGGAAGCACTGGAAGT
ZEP	GCTTGGAGGCAGAGTATTGAC	GCAGCCATTCTAGCCATTCC
CYP97C1	GATGAGAGCACAACAAGAGGTT	CGCCTTATTAACACTGGTGGAT

化合物 Compound	分子式 Formula	离子模式 ESM	质荷比 m/z	保留时间 Retention/min	二级鉴定分数 MS2 score
吲哚-3-羧酸 Indole-3-carboxylic acid	C₉H₇NO₂	-	160.04	1.84	43.60
吲哚 Indole	C₈H₇N	-	152.01	1.88	63.40
吲哚-5-羧酸 Indole-5-carboxylic acid	C₉H₇NO₂	-	160.04	2.05	33.60
3-甲基羟基吲哚 3-Methyldioxyindole	C₉H₉NO₂	-	162.06	2.27	38.70
5,6-二羟基吲哚 5,6-Dihydroxyindole	C₈H₇NO₂	-	297.09	2.67	27.00
吲哚-3-甲醛 Indole-3-carboxaldehyde	C₉H₇N_O	+	110.04	3.88	0.00
5,6-吲哚醌-2-羧酸 5,6-Indolequinone-2-carboxylic acid	C₉H₅NO₄	+	383.05	1.81	22.00
1-萘乙酸乙酯 Ethyl 1-naphthylacetic acid	C₁₄H₁₄O₂	+	215.11	3.60	34.30
赤霉素3 Gibberellin A3	C₁₉H₂₂O₆	-	391.15	3.16	74.30
赤霉素4 Gibberellin A4	C₁₉H₂₄O₅	-	663.29	6.78	78.10
赤霉素36 Gibberellin A36	C₂₀H₂₆O₆	+	345.17	7.76	66.80
邻β-d-葡糖基玉米素 O-Beta-D-glucosylzeatin	C₁₆H₂₃N₅O₆	-	402.13	3.65	74.30
油菜素内酯 Brassinolide	C₂₈H₄₈O₆	+	445.34	10.88	53.40
二氢茉莉酸 Dihydrojasmonic acid	C₁₂H₂₀O₃	-	233.12	4.42	36.50
（-）-茉莉酸（-）-Jasmonic acid	C₁₂H₁₈O₃	+	175.11	4.99	66.60
茉莉酸 Jasmonic acid	C₁₂H₁₈O₃	+	175.11	7.34	69.00
5-甲氧基水杨酸 5-Methoxysalicylic acid	C₈H₈O₄	-	167.04	1.35	86.90
赤霉素20 Gibberellin A20	C₁₉H₂₄O₅	-	353.15	7.05	47.80
赤霉素8 Gibberellin A8	C₁₉H₂₄O₇	+	387.13	4.17	66.10
赤霉素19 Gibberellin A19	C₂₀H₂₆O₆	+	385.17	6.84	70.20
赤霉素9 Gibberellin A9	C₁₉H₂₄O₄	+	317.18	8.95	42.20
（+）-脱落酸（+）-Abscisic acid	C₁₅H₂₀O₄	+	247.13	5.70	81.20
油菜素甾酮 Castasterone	C₂₈H₄₈O₅	+	447.34	10.77	60.70
褪黑素 Melatonin	C₁₃H₁₆N₂O₂	-	277.12	2.80	29.50
对水杨酸 P-salicylic acid	C₇H₆O₃	-	137.03	1.56	28.90
茉莉酸甲酯 Methyl jasmonate	C₁₃H₂₀O₃	+	207.14	3.00	45.30
二氢茉莉酮酸甲酯 Methyl dihydrojasmonate	C₁₃H₂₂O₃	-	225.15	4.17	25.80
2-萘乙酸 2-Naphthylacetic acid	C₁₂H₁₀O₂	+	187.08	8.05	33.90
玉米素 Zeatin	C₁₀H₁₃N₅O	+	220.12	1.57	26.50
玉米素核苷 cis-Zeatin riboside	C₁₅H₂₁N₅O₅	+	374.15	3.43	78.90
3-吲哚丙烯酸 3-Indoleacrylic acid	C₁₁H₉NO₂	-	208.03	3.07	43.60

化合物 Compound	分子式 Formula	离子模式 ESM	质荷比 m/z	保留时间 Retention/min	二级鉴定分数 MS2 score
吲哚-3-羧酸 Indole-3-carboxylic acid	C₉H₇NO₂	-	160.04	1.84	43.60
吲哚 Indole	C₈H₇N	-	152.01	1.88	63.40
吲哚-5-羧酸 Indole-5-carboxylic acid	C₉H₇NO₂	-	160.04	2.05	33.60
3-甲基羟基吲哚 3-Methyldioxyindole	C₉H₉NO₂	-	162.06	2.27	38.70
5,6-二羟基吲哚 5,6-Dihydroxyindole	C₈H₇NO₂	-	297.09	2.67	27.00
吲哚-3-甲醛 Indole-3-carboxaldehyde	C₉H₇N_O	+	110.04	3.88	0.00
5,6-吲哚醌-2-羧酸 5,6-Indolequinone-2-carboxylic acid	C₉H₅NO₄	+	383.05	1.81	22.00
1-萘乙酸乙酯 Ethyl 1-naphthylacetic acid	C₁₄H₁₄O₂	+	215.11	3.60	34.30
赤霉素3 Gibberellin A3	C₁₉H₂₂O₆	-	391.15	3.16	74.30
赤霉素4 Gibberellin A4	C₁₉H₂₄O₅	-	663.29	6.78	78.10
赤霉素36 Gibberellin A36	C₂₀H₂₆O₆	+	345.17	7.76	66.80
邻β-d-葡糖基玉米素 O-Beta-D-glucosylzeatin	C₁₆H₂₃N₅O₆	-	402.13	3.65	74.30
油菜素内酯 Brassinolide	C₂₈H₄₈O₆	+	445.34	10.88	53.40
二氢茉莉酸 Dihydrojasmonic acid	C₁₂H₂₀O₃	-	233.12	4.42	36.50
（-）-茉莉酸（-）-Jasmonic acid	C₁₂H₁₈O₃	+	175.11	4.99	66.60
茉莉酸 Jasmonic acid	C₁₂H₁₈O₃	+	175.11	7.34	69.00
5-甲氧基水杨酸 5-Methoxysalicylic acid	C₈H₈O₄	-	167.04	1.35	86.90
赤霉素20 Gibberellin A20	C₁₉H₂₄O₅	-	353.15	7.05	47.80
赤霉素8 Gibberellin A8	C₁₉H₂₄O₇	+	387.13	4.17	66.10
赤霉素19 Gibberellin A19	C₂₀H₂₆O₆	+	385.17	6.84	70.20
赤霉素9 Gibberellin A9	C₁₉H₂₄O₄	+	317.18	8.95	42.20
（+）-脱落酸（+）-Abscisic acid	C₁₅H₂₀O₄	+	247.13	5.70	81.20
油菜素甾酮 Castasterone	C₂₈H₄₈O₅	+	447.34	10.77	60.70
褪黑素 Melatonin	C₁₃H₁₆N₂O₂	-	277.12	2.80	29.50
对水杨酸 P-salicylic acid	C₇H₆O₃	-	137.03	1.56	28.90
茉莉酸甲酯 Methyl jasmonate	C₁₃H₂₀O₃	+	207.14	3.00	45.30
二氢茉莉酮酸甲酯 Methyl dihydrojasmonate	C₁₃H₂₂O₃	-	225.15	4.17	25.80
2-萘乙酸 2-Naphthylacetic acid	C₁₂H₁₀O₂	+	187.08	8.05	33.90
玉米素 Zeatin	C₁₀H₁₃N₅O	+	220.12	1.57	26.50
玉米素核苷 cis-Zeatin riboside	C₁₅H₂₁N₅O₅	+	374.15	3.43	78.90
3-吲哚丙烯酸 3-Indoleacrylic acid	C₁₁H₉NO₂	-	208.03	3.07	43.60

代谢酶Enzyme name	缩写Abbreviation	KO 编码KO ID	unigene数量Number of unigene
1-脱氧-D-木酮糖5-磷酸合酶1-Deoxy-D-xylulose 5-phosphate synthase	DXS	K01662	3
15-cis-phytoene合成酶15-cis-Phytoene synthase	PSY	K02291	1
15 - cis - Phytoene去饱和酶15-cis-Phytoene desaturase	PDS	K02293	1
前番茄红素异构酶Prolycopene isomerase	CRTISO	K09835	1
番茄红素β-环化酶Lycopene beta-cyclase	LCYB	K06443	1
番茄红素环化酶Lycopene epsilon-cyclase	LCYE	K06444	1
胡萝卜素羟基Beta-ring hydroxylase	CHYB	K15747	2
类胡萝卜素ε-羟化酶Carotenoid epsilon hydroxylase	CYP97C1	K09837	1
β -胡萝卜素异构酶Beta-carotene isomerase	DWARF27	K17911	3
内酯合成酶Carlactone synthase	CCD8	K17913	1
玉米黄质环氧化酶Zeaxanthin epoxidase	ZEP	K09838	3
紫黄质脱环氧化酶Violaxanthin de-epoxidase	VDE	K09839	1
9-顺式-环氧类胡萝卜素二加氧酶9-cis-Epoxycarotenoid dioxygenase	NCED	K09840	2
黄氧素脱氢酶Xanthoxin dehydrogenase	ABA2	K09841	3
（+）- 脱落酸8’羟化酶（+）-Abscisic acid 8’-hydroxylase	CYP707A	K09843	1
脱落酸β-葡萄糖基转移酶Abscisate beta-glucosyltransferase	AOG	K14595	1