信号分子H2O2调控糙皮侧耳菌盖颜色形成的机制初探

doi:10.16420/j.issn.0513-353x.2022-0546

摘要/Abstract

摘要：

通过外源喷施H₂O₂及其清除剂N, N'-二甲基硫脲（N, N'-Dimethylthiourea，DMTU）试验明确了H₂O₂能够参与调控糙皮侧耳（Pleurotus ostreatus）菌盖颜色的形成。利用已报道的双孢蘑菇（Agaricus bisporus）黑色素合成途径为参考，通过序列比对鉴定出28个参与黑色素合成途径的基因，并通过qPCR分析外源H₂O₂对不同途径中关键酶基因相对表达量的影响。结果表明H₂O₂信号分子一方面可调控不同黑色素合成途径中关键酶基因的表达，使得糙皮侧耳中不同类型黑色素比例发生变化而影响菌盖颜色；另一方面可促进铜转录因子基因mac、铜转运蛋白基因ctr及多酚氧化酶基因ppo1的高表达，进而影响菌盖颜色形成。

关键词: 糙皮侧耳, 菌盖颜色, 过氧化氢, 多酚氧化酶, 铜离子转运

Abstract:

By spraying H₂O₂ and its scavenger N，N'-dimethylthiourea（DMTU）externally，it was clarified that H₂O₂ can participate in regulating the formation of Pleurotus ostreatus cap color. Using the reported Agaricus bisporus as a reference，28 genes involved in the melanin synthesis pathways were identified through sequence alignment，and the effect of exogenous H₂O₂ on the relative expression of key enzyme genes in different pathways was analyzed by qPCR. The results showed that H₂O₂ signaling molecule can change the proportion of different types of melanin in P. ostreatus and affecting the color of the cap by regulating the expression of key enzyme genes in different melanin synthesis pathways. On the other hand，it can promote the high expression of copper transcription factor genes mac，copper transporter genes ctr and polyphenol oxidase gene ppo1，and finally affect the formation of cap color.

Key words: Pleurotus ostreatus, cap color, hydrogen peroxide, polyphenol oxidase, copper ion transport

中图分类号:

S646.141

侯潞丹, 颜克兴, 董帅, 刘星雨, 常明昌, 孟俊龙. 信号分子H₂O₂调控糙皮侧耳菌盖颜色形成的机制初探[J]. 园艺学报, 2023, 50(6): 1243-1254.

HOU Ludan, YAN Kexing, DONG Shuai, LIU Xingyu, CHANG Mingchang, MENG Junlong. Preliminary Study on the Mechanism of Signal Molecule H₂O₂ Regulating the Formation of Pleurotus ostreatus Cap Color[J]. Acta Horticulturae Sinica, 2023, 50(6): 1243-1254.

图/表 9

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引物	正向序列（5′-3′）	反向序列（5′-3′）
Primer	Forward sequence	Reverse sequence
qPCR_CM	GGGTAAATGGAGACGCAGAG	CTGGATTGGCTTGGAATTTGG
qPCR_4ABS	TGGGACGTACTCAGCAAAAG	ACACCAGTAACCAAAGACCC
qPCR_4ABH_1	CTCCCTACCTTCAGTGCATTC	TGAGGTGCTTGTTCTTTCCC
qPCR_4ABH_2	CTCCGAACATGACCCGAATAC	CTCACTAATCCCAAAGGCTCTC
qPCR_4ABH_3	GTTGTAGACTGTGAGAGGGAAG	TGAGACAAAGAATGGGAGTGG
qPCR_4ABH_4	GCAAGTCACATTCAAGGTCATC	TCCAGATTGCTATCGCTTCG
qPCR_GGT_1	CTGTCGAAAAGTTTGGTAGCG	CGTTGGGAGATGCATTTTGG
qPCR_GGT_2	GGTGGAGAACTATGACTTTCGG	TGCCGTGTCGTTCATATAAGC
qPCR_PDH	GTTTGCGACATCCCTGAAAAG	CGATGAACTCCGCTTCTACTG
qPCR_AT_1	CCTGATGCTTTCTACACCTCC	GTGAATGGAAAGGTTGTCGC
qPCR_AT_2	GGAGCGGTTGTGATATCTATAGG	TGAGGACATGAAGGATGGTTG
qPCR_ppo1	ACCCTTTGTTCTGGCTTCATC	TTCGTGATATTGTCCAGGCG
qPCR_ppo2	ACCCCAACCTTCAAAGTACAC	TGAGATCCATTGACTGCCAC
qPCR_ppo3	AAAGTGGGCTCCTAACGATC	GTAATGTTCTCGAAACGCTGG
qPCR_ppo4	GTCCAACGATCCCATTTTCTG	ATGAGCGGGAAAGGTATGAAG
qPCR_mac1	CAAAACAAAGGTGCCAGACAG	CATTCACCGCCAGTTTTACAC
qPCR_mac2	CCTATTGAACCAGCTCTTCCC	CATTGGCATCCACGAGTACT
qPCR_ctr1	TTTGCCATACAGTCCCTTCTC	CGATGAGATAGCCAGCATGG
qPCR_ctr2	TTACTCGCTGCCTTTCGG	ATCCTTTTCTCCCATCGACG
qPCR_ctr3	AACCACATCTTCGACTACCAC	AACCACATCTTCGACTACCAC
qPCR_ctr4	GGTTTTCATGACATACAACGCG	TGGGAGTTGAAGACGAAATGG
tubulin	AGGCTTTCTTGCATTGGTACACGC	TATTCGCCTTCTTCCTCATCGGCA

引物	正向序列（5′-3′）	反向序列（5′-3′）
Primer	Forward sequence	Reverse sequence
qPCR_CM	GGGTAAATGGAGACGCAGAG	CTGGATTGGCTTGGAATTTGG
qPCR_4ABS	TGGGACGTACTCAGCAAAAG	ACACCAGTAACCAAAGACCC
qPCR_4ABH_1	CTCCCTACCTTCAGTGCATTC	TGAGGTGCTTGTTCTTTCCC
qPCR_4ABH_2	CTCCGAACATGACCCGAATAC	CTCACTAATCCCAAAGGCTCTC
qPCR_4ABH_3	GTTGTAGACTGTGAGAGGGAAG	TGAGACAAAGAATGGGAGTGG
qPCR_4ABH_4	GCAAGTCACATTCAAGGTCATC	TCCAGATTGCTATCGCTTCG
qPCR_GGT_1	CTGTCGAAAAGTTTGGTAGCG	CGTTGGGAGATGCATTTTGG
qPCR_GGT_2	GGTGGAGAACTATGACTTTCGG	TGCCGTGTCGTTCATATAAGC
qPCR_PDH	GTTTGCGACATCCCTGAAAAG	CGATGAACTCCGCTTCTACTG
qPCR_AT_1	CCTGATGCTTTCTACACCTCC	GTGAATGGAAAGGTTGTCGC
qPCR_AT_2	GGAGCGGTTGTGATATCTATAGG	TGAGGACATGAAGGATGGTTG
qPCR_ppo1	ACCCTTTGTTCTGGCTTCATC	TTCGTGATATTGTCCAGGCG
qPCR_ppo2	ACCCCAACCTTCAAAGTACAC	TGAGATCCATTGACTGCCAC
qPCR_ppo3	AAAGTGGGCTCCTAACGATC	GTAATGTTCTCGAAACGCTGG
qPCR_ppo4	GTCCAACGATCCCATTTTCTG	ATGAGCGGGAAAGGTATGAAG
qPCR_mac1	CAAAACAAAGGTGCCAGACAG	CATTCACCGCCAGTTTTACAC
qPCR_mac2	CCTATTGAACCAGCTCTTCCC	CATTGGCATCCACGAGTACT
qPCR_ctr1	TTTGCCATACAGTCCCTTCTC	CGATGAGATAGCCAGCATGG
qPCR_ctr2	TTACTCGCTGCCTTTCGG	ATCCTTTTCTCCCATCGACG
qPCR_ctr3	AACCACATCTTCGACTACCAC	AACCACATCTTCGACTACCAC
qPCR_ctr4	GGTTTTCATGACATACAACGCG	TGGGAGTTGAAGACGAAATGG
tubulin	AGGCTTTCTTGCATTGGTACACGC	TATTCGCCTTCTTCCTCATCGGCA

关键酶编号 Key enzyme number	基因功能 Gene function	基因缩写 Gene abbreviation	基因编号 Gene number
Ⅰ	分支酸变位酶 Chorismate mutase	CM	g11035
Ⅱ	4-氨基苯甲酸合成酶 4-Aminobenzote synthase	4ABS	g1809
Ⅲ	4-氨基苯甲酸羟化酶1 4-Aminobenzote hydroxylase_1	4ABH_1	g2093
Ⅲ	4-氨基苯甲酸羟化酶2 4-Aminobenzote hydroxylase_2	4ABH_2	g3895
Ⅲ	4-氨基苯甲酸羟化酶3 4-Aminobenzote hydroxylase_3	4ABH_3	g12346
Ⅲ	4-氨基苯甲酸羟化酶4 4-Aminobenzote hydroxylase_4	4ABH_4	g9216
Ⅳ	γ-谷氨酰胺转移酶1 γ-Glutaminyltransferase_1	GGT_1	g649
Ⅳ	γ-谷氨酰胺转移酶2 γ-Glutaminyltransferase_2	GGT_2	g8358
Ⅴ	预苯甲酸脱水酶 Prephenate dehydratase	PDHtase	g9753
Ⅵ	预苯酸脱氢酶 Prephenate dehydrogenase	PDH	g10441
Ⅶ/Ⅷ	（4-羟基）苯基丙酮酸氨基转移酶1 (4-Hydroxy) phenylpyruvate aminoransferase_1	AT_1	g4186
Ⅶ/Ⅷ	（4-羟基）苯基丙酮酸氨基转移酶2 (4-Hydroxy) phenylpyruvate aminoransferase_2	AT_2	g7673
Ⅸ	多酚氧化酶1 Polyphenooxidase_1	PPO1	g7596
Ⅸ	多酚氧化酶2 Polyphenooxidase_2	PPO2	g3399
Ⅸ	多酚氧化酶3 Polyphenooxidase_3	PPO3	g3812
Ⅸ	多酚氧化酶4 Polyphenooxidase_4	PPO4	g11413
Ⅹ	苯丙氨酸解氨酶1 Phenylalnine ammonialyase_1	PAL1	g9868
Ⅹ	苯丙氨酸解氨酶2 Phenylalnine ammonialyase_2	PAL2	g9752
Ⅻ	4-香豆酸辅酶A连接酶1 4-Coumarate CoA ligase_1	4CL1	g5055
Ⅻ	4-香豆酸辅酶A连接酶2 4-Coumarate CoA ligase_2	4CL2	g1908
Ⅻ	4-香豆酸辅酶A连接酶3 4-Coumarate CoA ligase_3	4CL3	g7302
Ⅻ	4-香豆酸辅酶A连接酶4 4-Coumarate CoA ligase_4	4CL4	g8331
Ⅻ	4-香豆酸辅酶A连接酶5 4-Coumarate CoA ligase_5	4CL5	g11056
Ⅻ	4-香豆酸辅酶A连接酶6 4-Coumarate CoA ligase_6	4CL6	g1913
Ⅻ	4-香豆酸辅酶A连接酶7 4-Coumarate CoA ligase_7	4CL7	g5185
Ⅻ	4-香豆酸辅酶A连接酶8 4-Coumarate CoA ligase_8	4CL8	g333
Ⅻ	4-香豆酸辅酶A连接酶9 4-Coumarate CoA ligase_9	4CL9	g9792
Ⅻ	4-香豆酸辅酶A连接酶10 4-Coumarate CoA ligase_10	4CL10	g9146

关键酶编号 Key enzyme number	基因功能 Gene function	基因缩写 Gene abbreviation	基因编号 Gene number
Ⅰ	分支酸变位酶 Chorismate mutase	CM	g11035
Ⅱ	4-氨基苯甲酸合成酶 4-Aminobenzote synthase	4ABS	g1809
Ⅲ	4-氨基苯甲酸羟化酶1 4-Aminobenzote hydroxylase_1	4ABH_1	g2093
Ⅲ	4-氨基苯甲酸羟化酶2 4-Aminobenzote hydroxylase_2	4ABH_2	g3895
Ⅲ	4-氨基苯甲酸羟化酶3 4-Aminobenzote hydroxylase_3	4ABH_3	g12346
Ⅲ	4-氨基苯甲酸羟化酶4 4-Aminobenzote hydroxylase_4	4ABH_4	g9216
Ⅳ	γ-谷氨酰胺转移酶1 γ-Glutaminyltransferase_1	GGT_1	g649
Ⅳ	γ-谷氨酰胺转移酶2 γ-Glutaminyltransferase_2	GGT_2	g8358
Ⅴ	预苯甲酸脱水酶 Prephenate dehydratase	PDHtase	g9753
Ⅵ	预苯酸脱氢酶 Prephenate dehydrogenase	PDH	g10441
Ⅶ/Ⅷ	（4-羟基）苯基丙酮酸氨基转移酶1 (4-Hydroxy) phenylpyruvate aminoransferase_1	AT_1	g4186
Ⅶ/Ⅷ	（4-羟基）苯基丙酮酸氨基转移酶2 (4-Hydroxy) phenylpyruvate aminoransferase_2	AT_2	g7673
Ⅸ	多酚氧化酶1 Polyphenooxidase_1	PPO1	g7596
Ⅸ	多酚氧化酶2 Polyphenooxidase_2	PPO2	g3399
Ⅸ	多酚氧化酶3 Polyphenooxidase_3	PPO3	g3812
Ⅸ	多酚氧化酶4 Polyphenooxidase_4	PPO4	g11413
Ⅹ	苯丙氨酸解氨酶1 Phenylalnine ammonialyase_1	PAL1	g9868
Ⅹ	苯丙氨酸解氨酶2 Phenylalnine ammonialyase_2	PAL2	g9752
Ⅻ	4-香豆酸辅酶A连接酶1 4-Coumarate CoA ligase_1	4CL1	g5055
Ⅻ	4-香豆酸辅酶A连接酶2 4-Coumarate CoA ligase_2	4CL2	g1908
Ⅻ	4-香豆酸辅酶A连接酶3 4-Coumarate CoA ligase_3	4CL3	g7302
Ⅻ	4-香豆酸辅酶A连接酶4 4-Coumarate CoA ligase_4	4CL4	g8331
Ⅻ	4-香豆酸辅酶A连接酶5 4-Coumarate CoA ligase_5	4CL5	g11056
Ⅻ	4-香豆酸辅酶A连接酶6 4-Coumarate CoA ligase_6	4CL6	g1913
Ⅻ	4-香豆酸辅酶A连接酶7 4-Coumarate CoA ligase_7	4CL7	g5185
Ⅻ	4-香豆酸辅酶A连接酶8 4-Coumarate CoA ligase_8	4CL8	g333
Ⅻ	4-香豆酸辅酶A连接酶9 4-Coumarate CoA ligase_9	4CL9	g9792
Ⅻ	4-香豆酸辅酶A连接酶10 4-Coumarate CoA ligase_10	4CL10	g9146

信号分子H₂O₂调控糙皮侧耳菌盖颜色形成的机制初探

Preliminary Study on the Mechanism of Signal Molecule H₂O₂ Regulating the Formation of Pleurotus ostreatus Cap Color

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