番木瓜PG基因家族鉴定及其在果实软化中的功能

doi:10.13560/j.cnki.biotech.bull.1985.2025-0437

摘要/Abstract

摘要：

目的多聚半乳糖醛酸酶（PG）是调控果实软化的关键因子，鉴定番木瓜PG基因家族成员并进行功能分析，为揭示番木瓜采后软化的分子机制提供理论基础。方法基于‘紫晖’番木瓜基因组数据，全面鉴定CpPG家族成员，分析其蛋白理化性质、系统进化关系、保守基序、基因结构、基因复制事件、启动子顺式作用元件和表达水平；进一步利用果实瞬时过表达试验，探究CpPG1、CpPG2、CpPG28在番木瓜果实采后软化中的功能。结果在‘紫晖’番木瓜基因组中鉴定出38个PG家族成员，命名为CpPG1-CpPG38，这些成员不均匀分布在8条染色体上。系统进化分析表明CpPG成员聚类为亚组A-F，各亚组内基因成员的外显子/内含子结构相对保守，不同组别CpPG成员在保守结构域和蛋白基序具有显著差异。共线性分析结果表明，番木瓜PG家族存在基因复制事件和基因串联重复。CpPG成员的启动子区域包含多种光响应、植物生长发育与逆境胁迫、植物激素应答相关作用元件，参与多种生长发育进程。通过分析38个CpPG基因的表达模式，筛选出CpPG1、CpPG2和CpPG28 3个核心成员，这些基因在成熟番木瓜果实中高表达并响应乙烯诱导。瞬时过表达试验进一步证实，CpPG1、CpPG2和CpPG28均能显著降低果实硬度，揭示其在番木瓜果实软化中的关键作用。结论番木瓜基因组中鉴定出38个CpPG基因，挖掘3个调控番木瓜果实软化的关键家族成员CpPG1、CpPG2和CpPG28。

关键词: 番木瓜, 果实, 成熟软化, 多聚半乳糖醛酸酶, 基因家族, 表达模式, 系统进化, 功能分析

Abstract:

Objective Polygalacturonase (PG) plays a pivotal role in the regulation of fruit softening. This study is aimed to identify the PG gene family members in papaya (Carica papaya L.) and analyze their functions, providing a theoretical framework for understanding the molecular mechanisms behind postharvest softening in papaya. Method Based on the genome data of ‘Zihui’ papaya, the CpPG family members were systematically identified,and their protein properties, evolutionary relationships, conserved motifs, gene structures, gene duplication events, cis-acting elements, and expression patterns were thoroughly investigated. Additionally, transient overexpression experiments in papaya fruit were conducted to assess the roles of CpPG1, CpPG2, and CpPG28 in fruit softening. Result A total of 38 PG genes were identified in the ‘Zihui’ papaya genome, named as CpPG1 to CpPG38, and they were distributed unevenly across 8 chromosomes. Phylogenetic analysis grouped the CpPG genes into subgroup A-F, with relatively conserved exon/intron structures within each group. However, significant differences in conserved domains and protein motifs were observed among subgroups. Collinearity analysis revealed the evidence of gene duplication and tandem repeat events within the papaya CpPG family. Promoter regions of CpPG genes contained diverse cis-elements associated with light responsiveness, plant development, stress adaptation, and hormone signaling, indicating their involvement in various developmental processes. Expression pattern analysis identified three key genes, CpPG1, CpPG2, and CpPG28, which were highly expressed in ripe papaya fruit and responsive to ethylene. Transient overexpression experiments confirmed that these genes significantly decreased fruit firmness, highlighting their critical roles in fruit softening. Conclusion This study identified 38 CpPG genes in the papaya genome and highlighted CpPG1, CpPG2, and CpPG28 as key regulators of papaya fruit softening.

Key words: papaya, fruit, ripening and softening, polygalacturonase, gene family, gene expression pattern, evolutionary relationship, functional analysis

徐泽, 周陈平, 邝瑞彬, 吴夏明, 杨敏, 刘传和, 贺涵, 魏岳荣. 番木瓜PG基因家族鉴定及其在果实软化中的功能[J]. 生物技术通报, 2026, 42(3): 349-361.

XU Ze, ZHOU Chen-ping, KUANG Rui-bin, WU Xia-ming, YANG Min, LIU Chuan-he, HE Han, WEI Yue-rong. Identification of PG Gene Family and Their Roles in Papaya Fruit Softening[J]. Biotechnology Bulletin, 2026, 42(3): 349-361.

图/表 10

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引物名称 Gene name	引物序列 Primer sequence （5′‒3′）
CpPG1-F	AATGGTGGTGCGTATAGATGGAAC
CpPG1-R	GGGAGTTGAGAAACCCGATTGAC
CpPG2-F	CACCCTTCATTTCTCAGCTCCATT
CpPG2-R	TTTCTCCCAAGCCTCCTCAAATAC
CpPG28-F	ACCATTGACGGACAAGGCTCTAT
CpPG28-R	TCAGACAAGGTGAACTCGACAAGA
OE-CpPG1-F	CCAAATCGACTCTAGTCTAGAATGACGACAATCCGCTCTCACAAC
OE-CpPG1-R	GCTCACCATCTGCAGAAGCTTCAAGCAACTATTGGGCTGAACCG
OE-CpPG2-F	CCAAATCGACTCTAGTCTAGAATGGCTGTTCTATATTATGATAATTATAATCGTACCC
OE-CpPG2-R	GCTCACCATCTGCAGAAGCTTCCAGATTATCATCTTAAACATGACGAGCTT
OE-CpPG28-F	CCAAATCGACTCTAGTCTAGAATGAAGATGCCAGTAGCATTTTTCTTGC
OE-CpPG28-R	GCTCACCATCTGCAGAAGCTTTACCGGGACAGCCACAGTTTCA
EIF4A-F	AGGCAGGCAAGAGAAGAT
EIF4A-R	TTCATACCGAGTAGCGATTC

引物名称 Gene name	引物序列 Primer sequence （5′‒3′）
CpPG1-F	AATGGTGGTGCGTATAGATGGAAC
CpPG1-R	GGGAGTTGAGAAACCCGATTGAC
CpPG2-F	CACCCTTCATTTCTCAGCTCCATT
CpPG2-R	TTTCTCCCAAGCCTCCTCAAATAC
CpPG28-F	ACCATTGACGGACAAGGCTCTAT
CpPG28-R	TCAGACAAGGTGAACTCGACAAGA
OE-CpPG1-F	CCAAATCGACTCTAGTCTAGAATGACGACAATCCGCTCTCACAAC
OE-CpPG1-R	GCTCACCATCTGCAGAAGCTTCAAGCAACTATTGGGCTGAACCG
OE-CpPG2-F	CCAAATCGACTCTAGTCTAGAATGGCTGTTCTATATTATGATAATTATAATCGTACCC
OE-CpPG2-R	GCTCACCATCTGCAGAAGCTTCCAGATTATCATCTTAAACATGACGAGCTT
OE-CpPG28-F	CCAAATCGACTCTAGTCTAGAATGAAGATGCCAGTAGCATTTTTCTTGC
OE-CpPG28-R	GCTCACCATCTGCAGAAGCTTTACCGGGACAGCCACAGTTTCA
EIF4A-F	AGGCAGGCAAGAGAAGAT
EIF4A-R	TTCATACCGAGTAGCGATTC

基因名称	基因号	氨基酸数	分子量	等电点	不稳定指数	脂肪系数	亲疏水性	信号肽
Gene name	Gene ID	Number of amino acids （aa）	Molecular weight （kD）	pI	Instability index	Aliphatic index	Hydropathicity index	Signal peptide
CpPG1	Cp_zihui11610	359	38.0	9.24	31.42	85.46	-0.060	+
CpPG2	Cp_zihui13876	437	48.2	5.69	34.70	77.12	-0.509	-
CpPG3	Cp_zihui12587	428	47.2	8.75	28.24	79.88	-0.373	+
CpPG4	Cp_zihui01876	390	42.2	8.63	44.78	87.95	-0.046	+
CpPG5	Cp_zihui19603	482	52.9	8.34	41.88	86.72	-0.111	-
CpPG6	Cp_zihui08791	447	48.7	7.57	36.53	89.80	-0.009	+
CpPG7	Cp_zihui00474	401	43.2	7.41	26.91	86.28	-0.026	+
CpPG8	Cp_zihui00470	278	30.7	8.94	28.18	79.93	-0.251	-
CpPG9	Cp_zihui00483	461	50.0	8.34	33.67	81.58	-0.211	-
CpPG10	Cp_zihui02232	458	49.0	6.30	44.43	77.86	-0.171	+
CpPG11	Cp_zihui06380	262	29.1	5.77	58.76	84.43	-0.274	-
CpPG12	Cp_zihui07099	432	45.8	5.35	45.09	77.59	-0.076	+
CpPG13	Cp_zihui08226	374	40.8	8.64	33.01	85.78	-0.102	+
CpPG14	Cp_zihui08889	488	53.6	5.38	35.37	90.20	-0.143	+
CpPG15	Cp_zihui09177	481	52.4	8.34	34.35	88.90	-0.025	+
CpPG16	Cp_zihui10261	349	37.8	7.89	35.97	86.07	-0.200	-
CpPG17	Cp_zihui10262	371	40.4	8.71	30.26	84.88	-0.188	-
CpPG18	Cp_zihui10270	335	35.8	6.72	45.96	94.84	-0.067	+
CpPG19	Cp_zihui10273	431	47.5	8.55	44.22	88.19	-0.016	+
CpPG20	Cp_zihui10665	475	51.6	5.48	50.72	83.28	-0.080	-
CpPG21	Cp_zihui10984	498	54.8	5.12	37.87	89.22	-0.144	-
CpPG22	Cp_zihui11565	530	58.3	5.71	31.68	87.19	0.085	-
CpPG23	Cp_zihui11577	425	45.4	6.78	29.06	84.24	-0.052	+
CpPG24	Cp_zihui11602	356	37.6	9.6	28.92	87.36	0.043	+
CpPG25	Cp_zihui11601	393	42.0	9.68	36.76	80.87	-0.097	+
CpPG26	Cp_zihui11608	395	43.1	7.58	38.66	84.89	-0.072	+
CpPG27	Cp_zihui11609	403	42.8	5.40	38.28	80.77	-0.082	+
CpPG28	Cp_zihui11883	457	49.6	5.59	48.03	90.83	-0.042	+
CpPG29	Cp_zihui12006	494	53.1	4.80	53.12	73.00	-0.252	+
CpPG30	Cp_zihui13178	477	52.2	5.35	33.45	85.97	-0.203	+
CpPG31	Cp_zihui13181	454	49.9	8.72	43.42	88.41	-0.148	+
CpPG32	Cp_zihui13193	499	54.3	5.20	34.53	81.42	-0.164	+
CpPG33	Cp_zihui14064	477	52.8	6.71	44.22	88.68	-0.208	-
CpPG34	Cp_zihui14065	429	46.3	5.38	42.46	91.17	-0.078	+
CpPG35	Cp_zihui16353	240	27.3	6.11	29.48	116.88	0.202	+
CpPG36	Cp_zihui16358	277	30.2	7.21	42.86	92.82	-0.201	-
CpPG37	Cp_zihui16904	536	60.3	8.70	44.09	87.95	-0.238	-
CpPG38	Cp_zihui19689	415	45.0	4.61	39.69	77.49	-0.295	+

基因名称	基因号	氨基酸数	分子量	等电点	不稳定指数	脂肪系数	亲疏水性	信号肽
Gene name	Gene ID	Number of amino acids （aa）	Molecular weight （kD）	pI	Instability index	Aliphatic index	Hydropathicity index	Signal peptide
CpPG1	Cp_zihui11610	359	38.0	9.24	31.42	85.46	-0.060	+
CpPG2	Cp_zihui13876	437	48.2	5.69	34.70	77.12	-0.509	-
CpPG3	Cp_zihui12587	428	47.2	8.75	28.24	79.88	-0.373	+
CpPG4	Cp_zihui01876	390	42.2	8.63	44.78	87.95	-0.046	+
CpPG5	Cp_zihui19603	482	52.9	8.34	41.88	86.72	-0.111	-
CpPG6	Cp_zihui08791	447	48.7	7.57	36.53	89.80	-0.009	+
CpPG7	Cp_zihui00474	401	43.2	7.41	26.91	86.28	-0.026	+
CpPG8	Cp_zihui00470	278	30.7	8.94	28.18	79.93	-0.251	-
CpPG9	Cp_zihui00483	461	50.0	8.34	33.67	81.58	-0.211	-
CpPG10	Cp_zihui02232	458	49.0	6.30	44.43	77.86	-0.171	+
CpPG11	Cp_zihui06380	262	29.1	5.77	58.76	84.43	-0.274	-
CpPG12	Cp_zihui07099	432	45.8	5.35	45.09	77.59	-0.076	+
CpPG13	Cp_zihui08226	374	40.8	8.64	33.01	85.78	-0.102	+
CpPG14	Cp_zihui08889	488	53.6	5.38	35.37	90.20	-0.143	+
CpPG15	Cp_zihui09177	481	52.4	8.34	34.35	88.90	-0.025	+
CpPG16	Cp_zihui10261	349	37.8	7.89	35.97	86.07	-0.200	-
CpPG17	Cp_zihui10262	371	40.4	8.71	30.26	84.88	-0.188	-
CpPG18	Cp_zihui10270	335	35.8	6.72	45.96	94.84	-0.067	+
CpPG19	Cp_zihui10273	431	47.5	8.55	44.22	88.19	-0.016	+
CpPG20	Cp_zihui10665	475	51.6	5.48	50.72	83.28	-0.080	-
CpPG21	Cp_zihui10984	498	54.8	5.12	37.87	89.22	-0.144	-
CpPG22	Cp_zihui11565	530	58.3	5.71	31.68	87.19	0.085	-
CpPG23	Cp_zihui11577	425	45.4	6.78	29.06	84.24	-0.052	+
CpPG24	Cp_zihui11602	356	37.6	9.6	28.92	87.36	0.043	+
CpPG25	Cp_zihui11601	393	42.0	9.68	36.76	80.87	-0.097	+
CpPG26	Cp_zihui11608	395	43.1	7.58	38.66	84.89	-0.072	+
CpPG27	Cp_zihui11609	403	42.8	5.40	38.28	80.77	-0.082	+
CpPG28	Cp_zihui11883	457	49.6	5.59	48.03	90.83	-0.042	+
CpPG29	Cp_zihui12006	494	53.1	4.80	53.12	73.00	-0.252	+
CpPG30	Cp_zihui13178	477	52.2	5.35	33.45	85.97	-0.203	+
CpPG31	Cp_zihui13181	454	49.9	8.72	43.42	88.41	-0.148	+
CpPG32	Cp_zihui13193	499	54.3	5.20	34.53	81.42	-0.164	+
CpPG33	Cp_zihui14064	477	52.8	6.71	44.22	88.68	-0.208	-
CpPG34	Cp_zihui14065	429	46.3	5.38	42.46	91.17	-0.078	+
CpPG35	Cp_zihui16353	240	27.3	6.11	29.48	116.88	0.202	+
CpPG36	Cp_zihui16358	277	30.2	7.21	42.86	92.82	-0.201	-
CpPG37	Cp_zihui16904	536	60.3	8.70	44.09	87.95	-0.238	-
CpPG38	Cp_zihui19689	415	45.0	4.61	39.69	77.49	-0.295	+