贮藏温度影响泰国青柚采后果皮失绿的比较转录组学分析

doi:10.16420/j.issn.0513-353x.2025-0194

园艺学报 ›› 2026, Vol. 53 ›› Issue (5): 1253-1273.doi: 10.16420/j.issn.0513-353x.2025-0194

• 遗传育种·种质资源·分子生物学 • 上一篇下一篇

贮藏温度影响泰国青柚采后果皮失绿的比较转录组学分析

王斌¹^,², 曹秋艳¹^,^*(), 袁晓¹^,³, 朱云娜¹^,², 黄常健¹, 刘堂茂¹^,^*()

¹ 韶关学院生物与农业学院，广东省粤北食药资源利用与保护重点实验室，广东韶关 512005
² 粤北特色果蔬工程技术研究中心，韶关学院园艺工程技术研究中心，广东韶关 512005
³ 华南农业大学园艺学院，广东省果蔬保鲜重点实验室，广州 510642

收稿日期:2025-12-04 修回日期:2026-02-13 出版日期:2026-05-25 发布日期:2026-05-26
通讯作者:
^* E-mail：cqiuyan@163.com，
843249682@qq.com
基金资助:
2023年度韶关市科技支撑乡村振兴项目(230612178030335)

Comparative Transcriptomic Analysis of the Effects of Storage Temperature on Peel De-greening of Postharvest Thai Green Pomelo

WANG Bin¹^,², CAO Qiuyan¹^,^*(), YUAN Xiao¹^,³, ZHU Yunna¹^,², HUANG Changjian¹, LIU Tangmao¹^,^*()

¹ Guangdong Provincial Key Laboratory of Utilization and Conservation of Food and Medicinal Resources in Northern Region， College of Biology and Agriculture，Shaoguan University，Shaoguan， Guangdong 512005， China
² Guangdong Provincial Engineering and Technology Research Center of Special Fruit and Vegetables in Northern Region， Engineering and Technology Research Center of Shaoguan Horticulture in Shaoguan University，Shaoguan， Guangdong 512005， China
³ Guangdong Province Key Laboratory of Postharvest Science of Fruits and Vegetables， College of Horticulture，South China Agricultural University， Guangzhou 510642， China

Received:2025-12-04 Revised:2026-02-13 Published:2026-05-25 Online:2026-05-26
Contact:
^* E-mail：cqiuyan@163.com，
843249682@qq.com

摘要/Abstract

摘要：

果皮色泽是影响泰国青柚（Citrus maxima）商品性的重要指标，但贮藏温度是否以及如何影响泰国青柚果皮色泽仍不清楚。将八成熟的泰国青柚果实在5、15和25 ℃下贮藏15 d，利用比较转录组学方法结合表型和生理数据，探究贮藏温度对果皮色泽的影响。结果显示，5 ℃下贮藏能延缓果皮失绿，而15和25 ℃贮藏期间果皮快速失绿。在5、15和25 ℃下贮藏15 d，果皮总叶绿素含量分别下降了21.93%、64.53%和86.74%，总黄酮含量分别增加了8.56%、27.03%和35.46%。与贮藏前（0 d）相比，5和15℃贮藏5 d产生的差异表达基因（differentially expressed genes，DEG）均显著富集在植物激素信号转导等通路中，而25 ℃贮藏5 d产生的DEG显著富集在碳代谢等通路中。与5 ℃贮藏5 d相比，15或25 ℃贮藏所产生的DEG显著富集在卟啉与叶绿素代谢、光合作用天线蛋白、苯丙烷素生物合成和异黄酮生物合成等色素合成或代谢相关途径中；上调表达的DEG也显著富集在苯丙烷素生物合成和卟啉与叶绿素代谢途径中。通过加权基因共表达网络分析发现，洋红色（magenta）模块和黄色（yellow）模块内的基因与总叶绿素含量显著负相关，而棕色（brown）模块与总叶绿素含量显著正相关。此外，还鉴定到了119个与果皮失绿进程密切相关的转录因子基因，其中MYB、ERF和bHLH家族数量最多，且与叶绿素降解相关基因的表达模式高度相关。qRT-PCR分析表明，15和25 ℃贮藏前期叶绿素代谢和黄酮合成相关基因的表达显著上调。综上，贮藏温度显著影响泰国青柚果皮失绿，5 ℃贮藏能有效延缓果皮失绿，而15和25 ℃贮藏加速果皮失绿。15和25 ℃贮藏通过影响叶绿素代谢和和黄酮合成相关基因的表达促进果皮失绿，受贮藏温度诱导的MYB、ERF和bHLH可能参与泰国青柚果皮失绿的调控。

关键词: 柚, 采后, 贮藏温度, 果皮失绿, 转录组学分析

Abstract:

The peel color is a critical indicator influencing the commercial value of Thai green pomelo（Citrus maxima）fruit. However，the effects of storage temperature on peel color alterations remain unclear. In this study，fruit at 80% maturity were stored at 5，15，and 25 ℃ for 15 d to investigate temperature-dependent peel color changes through comparative transcriptomic analyses integrated with phenotypic and physiological data. The results showed that storage temperature significantly affected the peel color of Thai green pomelos. Storage at 5 ℃ delayed the de-greening，whereas rapid changes in de-greening were occurred under 15 and 25 ℃ conditions. After 15 d of storage，the total chlorophyll contents respectively decreased by 21.93%，64.53% and 86.74% at 5，15 and 25 ℃，while total flavonoid contents increased by 8.56%，27.03% and 35.46%，respectively. Compared to pre-storage（0 d），the differentially expressed genes（DEGs）identified after 5 d of storage at 5 and 15 ℃ were significantly enriched in several pathways，plant hormone signal transduction，while DEGs from storage at 25 ℃ were significantly enriched in pathways carbon metabolism. After 5 d of storage，DEGs generated at 15 or 25 ℃ compared to those at 5 ℃ were significantly enriched in pigment biosynthesis or metabolism-related pathways，such as porphyrin and chlorophyll metabolism，photosynthetic antenna proteins，the phenylpropanoid biosynthesis，and flavonoid biosynthesis. DEGs that were upregulated were also significantly enriched in the phenylpropanoid biosynthesis and porphyrin and chlorophyll metabolism pathways，weighted gene co-expression network analysis revealed that genes in the magenta and yellow modules were negatively correlated with total chlorophyll content，while that in the brown module showed a significant positive correlation with total chlorophyll content. Furthermore，119 transcription factor genes that closely related to the process of peel de-greening were identified，with the highest numbers belonging to the MYB，ERF，and bHLH families. Additionally，strong correlations were observed between the expression of MYB，ERF，and bHLH transcription factor genes and genes related to chlorophyll degradation. qRT-PCR analysis indicated that storage at 15 and 25 ℃ significantly upregulated the expression of genes related to chlorophyll metabolism and flavonoid biosynthesis during the early storage period. In conclusion，storage temperature significantly affects the de-greening of Thai green pomelo peels. Storage at 5 ℃ effectively delays the de-greening，while storage at 15 and 25 ℃ accelerates that progress. 15 and 25 ℃ storage facilitated the de-greening by influencing the expression of genes associated with chlorophyll metabolism and flavonoid biosynthesis，possibly involving MYB，ERF，and bHLH transcription factors induced by these temperatures in Thai green pomelos.

Key words: pomelo, postharvest, storage temperature, peel de-greening, transcriptomic analysis

王斌, 曹秋艳, 袁晓, 朱云娜, 黄常健, 刘堂茂. 贮藏温度影响泰国青柚采后果皮失绿的比较转录组学分析[J]. 园艺学报, 2026, 53(5): 1253-1273.

WANG Bin, CAO Qiuyan, YUAN Xiao, ZHU Yunna, HUANG Changjian, LIU Tangmao. Comparative Transcriptomic Analysis of the Effects of Storage Temperature on Peel De-greening of Postharvest Thai Green Pomelo[J]. Acta Horticulturae Sinica, 2026, 53(5): 1253-1273.

图/表 16

表1 qRT-PCR分析引物信息

Table 1 Primer information for qRT-PCR analysis

基因ID或NCBI登录号 Gene ID or NCBI accession No.	正向序列（5′-3′） Forward sequence	反向序列（5′-3′） Reverse sequence
GQ389668.1	TTCTCACTGAGGCTCCCCTT	AGAGAGAAAGCACAGCCTGG
Cg4g006180	AAGTGCGTTTGCGATTTGCT	AGCTGACCTTGGTACCGTTG
Cg5g014530	GATGGAAGACCCACCCCAAG	TCACCATTCCTGGCGACAAA
Cg7g023650	CCACCTCGCCTTCTGTTTCT	CAACACGGTCCTCTTCCCTC
Cg1g006040	AAGCCAGGTTCTGAGAGCAC	TCACTGCTGATGTGTCGGTC
Cg4g023550	GCTCAAGAAACCCCCAGACA	GGGTAATTTGGCCGTCGGTA
Cg6g013340	ACGTATGAGGCAAGGTGTGG	TCGTGGGCAGAAACCTCTTC
Cg1g011620	CAGTGGAGGTGGTGGTCAAG	AATGACTGGCCTCCATTGGC
Cg7g010980	TGGAATCCTTGTGTCTCGGC	TGATCCAGGGGGAAGGGAAT
Cg9g001040	CTCCTCTCGCCCTCCATCTA	CAGGAACGAAAACGGAACGG
Cg6g016490	TGGTGATGAAGTTGGGTCCG	TCCAGCATAGCCCTCGAAAC
Cg8g002960	ATGACTCCACAAGAGGCTGC	TCTGCTACATCTGCCTCCCT
Cg9g000740	CCGAGTTCAAAAGCACGCAA	GCCGAGGCTTTATTGGCTTG
Cg1g019920	ATGGCATCTCTCCAAGCGAC	GCTTTTTCCGGCGAATGGTT
Cg5g037790	CAACTCACCCTTCGCTGCTA	TGGTGTTCATCGGAGAACGG
Cg7g011250	GCATCCCGTCCGGTTTCTAT	ATAACTCATGCTGCTGCGGT

图1 泰国青柚果实在不同贮藏温度下的果皮色度和色素含量变化不同小写字母表示相同时期不同处理间差异显著（P < 0.05），下同。照片拍摄于贮藏15 d

Fig. 1 Changes in peel color and pigment contents of Thai green pomelo fruit under different storage temperatures Different lowercase letters in the same time indicate significant differences among different treatments at 0.05 level. The same below. Picture was captured at 15 d of storage

图2 RNA-Seq样品基因表达分析 A：主成分分析；B FPKM密度分布图；C：箱线图

Fig. 2 Analysis of gene expression in RNA-Seq samples A：Principal component analysis（PCA）；B：FPKM density distribution；C：Box plot

表2 测序数据概况

Table 2 Overview of sequencing data

样品编号 Sample No.	过滤后读数 Clean reads	过滤后碱基 Clean bases	GC含量/% GC content	Q30/%	与参考基因组比对率/% Mapping rate with reference genome
0 d-1	20 564 027	6 146 144 541	43.49	94.96	94.17
0 d-2	27 004 938	8 079 620 406	43.41	95.30	94.63
0 d-3	23 629 673	7 067 692 753	43.44	95.21	94.37
5 ℃-5 d-1	24 140 969	7 220 605 293	43.27	95.12	94.20
5 ℃-5 d-2	24 681 155	7 378 754 202	43.35	95.37	94.21
5 ℃-5 d-3	21 490 523	6 425 191 037	43.31	95.26	93.69
15 ℃-5 d-1	22 867 564	6 831 377 959	43.78	95.51	95.51
15 ℃-5 d-2	25 270 981	7 554 985 484	43.59	95.57	95.49
15 ℃-5 d-3	26 875 686	8 030 082 353	43.70	95.58	95.35
25 ℃-5 d-1	24 902 945	7 450 669 476	43.52	95.20	92.84
25 ℃-5 d-2	19 951 247	5 967 907 384	43.55	94.24	92.64
25 ℃-5 d-3	25 254 884	7 551 251 030	43.53	95.11	92.62

图3 泰国青柚不同温度下贮藏前后比较获得的差异表达基因的KEGG富集分析

Fig. 3 KEGG enrichment analysis of differentially expressed genes obtained from comparisons before and after storage of Thai green pomelo under different storage temperatures

图4 泰国青柚贮藏5 d时不同贮藏温度间差异表达基因的KEGG富集分析

Fig. 4 KEGG enrichment analysis of differentially expressed genes among Thai green pomelo stored at different temperatures for 5 d

图5 与果皮色度密切相关的基因模块及基因富集分析 A：分层聚类树；B：模块与性状的相关性；C ~ E：分别为洋红色、褐色和黄色模块中的基因功能富集

Fig. 5 Gene modules closely related to peel color and gene enrichment analysis A：Dynamic cluster tree；B：The correlation between modules and traits；C-E：Gene functional enrichment in magenta，brown and yellow module，respectively

图6 15和25 ℃上调表达的差异表达基因功能富集分析 A：5 ℃-5 d vs. 15 ℃-5 d比较组上调表达DEG的功能富集；B：5 ℃-5 d vs. 25 ℃-5 d比较组上调表达DEG的功能富集；C：两个比较组重叠DEG的功能富集

Fig. 6 Functional enrichment analysis of DEGs upregulated by storage at 15 ℃ and 25 ℃ A：Functional enrichment of upregulated DEGs from 5 ℃-5 d vs. 15 ℃-5 d comparison group；B：Functional enrichment of upregulated DEGs from 5 ℃-5 d vs. 25 ℃-5 d comparison group；C：Functional enrichment of overlapping DEGs in two comparison groups

图7 与果皮失绿密切相关的转录因子鉴定 A：与果皮失绿正相关的差异表达基因集；B：基因集KEGG富集分析；C：不同家族转录因子数量

Fig. 7 Identification of transcription factors closely related to fruit peel de-greening A：A set of differentially expressed genes positively correlated with fruit peel de-greening；B：KEGG enrichment analysis of gene set；C：The number of transcription factors in different families

图8 与果皮失绿密切相关的代表性转录因子的bHLH、MYB和ERF基因的表达模式 A ~ C：3个代表性的bHLH基因的表达模式；D ~ F：3个代表性的MYB基因的表达模式；G ~ I：3个代表性的ERF基因的表达模式

Fig. 8 The expression patterns of representative bHLH，MYB and ERF transcription factors gene closely related to fruit peel de-greening A-C：The expression patterns of three representative bHLH genes；D-F：The expression patterns of three representative MYB genes；G-I：Expression patterns of three representative ERF genes

图9 叶绿素降解途径相关基因的表达热图

Fig. 9 Expression heat-map of DEGs in the chlorophyll degradation pathway

图10 3个代表性叶绿素降解基因的表达模式

Fig. 10 Expression patterns of 3 representative chlorophyll degradation related genes

图11 MYB（A）、bHLH（B）和ERF（C）与叶绿素降解基因的网络图

Fig. 11 Network between MYB（A），bHLH（B），ERF（C）and chlorophyll degradation related genes

图12 黄酮合成相关基因的表达热图

Fig. 12 Expression heat-map of DEGs in the flavonoid biosynthesis pathway

图13 3个代表性黄酮合成相关基因的表达模式

Fig. 13 Expression patterns of 3 representative flavonoid biosynthesis related genes

图14 类胡萝卜素合成途径差异表达基因的表达热图

Fig. 14 Heatmap of DEGs in the carotenoid biosynthesis pathway

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贮藏温度影响泰国青柚采后果皮失绿的比较转录组学分析

Comparative Transcriptomic Analysis of the Effects of Storage Temperature on Peel De-greening of Postharvest Thai Green Pomelo

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贮藏温度影响泰国青柚采后果皮失绿的比较转录组学分析

Comparative Transcriptomic Analysis of the Effects of Storage Temperature on Peel De-greening of Postharvest Thai Green Pomelo

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