Functional Analysis of Pepper U-Box Domain-Containing Protein Gene CaPUB18 in Heat Tolerance

doi:10.16420/j.issn.0513-353x.2024-0897

Acta Horticulturae Sinica ›› 2026, Vol. 53 ›› Issue (4): 1175-1188.doi: 10.16420/j.issn.0513-353x.2024-0897

• Genetic & Breeding · Germplasm Resources · Molecular Biology • Previous Articles Next Articles

Functional Analysis of Pepper U-Box Domain-Containing Protein Gene CaPUB18 in Heat Tolerance

LI Ying¹, GUO Yuling¹, ZHAO Yue¹, YANG Qiaomin¹, CHEN Tao¹, AN Yuchang¹, MAO Zhangliang², LU Minghui¹^,^*()

¹ College of Horticulture，Northwest Agriculture and Forestry University，Yangling，Shaanxi 712100，China
² Xingping Agro-Products Quality and Safety Inspection and Testing Station，Xingping，Shaanxi 713100，China

Received:2025-10-13 Revised:2025-12-18 Online:2026-04-25 Published:2026-04-20
Contact: LU Minghui

Abstract

Abstract:

U-box E3 ubiquitin ligase（PUB）plays important roles in plant stress resistance，while the function of CaPUB18 gene in pepper heat tolerance is still unclear. Pepper CaPUB18 protein contains one U-box domain and four ARM domains，and presents closer phylogenetic relationship to its homologues from potato and tomato. CaPUB18 is localized in the cell membrane，and expressed in various tissues. During heat stress，its expression level increases initially，and then decreases. The silencing and heterologous overexpression of CaPUB18 compromised the heat tolerance of pepper and Arabidopsis，respectively，as evidenced by the aggravated leaf wilting，increased relative conductivity and malondialdehyde（MDA）content，accumulation of dead cells and H₂O₂，reduced total chlorophyll content，and inhibited induction of heat-tolerance marker genes. These results indicate that both silencing and overexpression of CaPUB18 are detrimental to the heat tolerance of pepper. This study highlights the importance of maintaining the stability of U-box protein gene CaPUB18 expression level to the formationof heat tolerance of pepper.

Key words: pepper, U-box containing protein, E3 ubiquitin ligase, CaPUB18, heat tolerance

LI Ying, GUO Yuling, ZHAO Yue, YANG Qiaomin, CHEN Tao, AN Yuchang, MAO Zhangliang, LU Minghui. Functional Analysis of Pepper U-Box Domain-Containing Protein Gene CaPUB18 in Heat Tolerance[J]. Acta Horticulturae Sinica, 2026, 53(4): 1175-1188.

Figures/Tables 10

Table 1 Primers and sequence used in this study

引物名称 Primer name	序列（5′-3′） Sequence	用途 Purpose
pART27-GFP-CaPUB18-F	GATGAACTATACAAAGAATTCATGATCAACGACAAATCTAACGA	基因CDS区全长扩增 Amplification of entire CDS（Coding DNA sequence）
pART27-GFP-CaPUB18-R	CAGGACTCTAGATTAGGTACCTTACCACACATTAATGAATTGATCC	基因CDS区全长扩增 Amplification of entire CDS（Coding DNA sequence）
pTRV2- CaPUB18-F	GCTCTAGATCGAAACTCGATACGTTTGATTG	基因沉默特异片段扩增 Construction of gene-silencing vector
pTRV2- CaPUB18-R	CGGGATCCTCCACCACTTCTTTAACTTCCAC	基因沉默特异片段扩增 Construction of gene-silencing vector
qCaPUB18-F	GCTTTTGGGCACACTTGAGG	基因表达分析 Analysis of gene expression
qCaPUB18-R	TGTGTGATTCTCGGGTGGTG
qCaUbi3-F	TGTCCATCTGCTCTCTGTTG
qCaUbi3-R	CACCCCAAGCACAATAAGAC
qCaHSP70.1-F	CAGGTGTGCTAGTTCAGGTGT
qCaHSP70.1-R	TGACCTGAGGCACTCCTCTT
qCaHsfA2-F	GTAGCATCAGTAGCCACAGC
qCaHsfA2-R	CAAGCAACTCTTCCCAAATA
AtActin2-F	CGCTCTTTCTTTCCAAGCTCAT
AtActin2-R	CAAATCCAGCCTTCACCAT

Fig. 1 Sequence characterization of CaPUB18 A：Analysis of conservative domains；B：Analysis of phylogenetic relationship

Table 2 Prediction cis-acting elements in CaPUB18 promoter

元件 Component	序列 Sequence	数量 Quantity	功能 Function
ABRE	ACGTG	2	响应脱落酸信号 Involved in the ABA-responsiveness
CGTCA-motif	CGTCA	1	响应茉莉酸甲酯信号 Involved in the MeJA-responsiveness
G-box	CACGTC	2	响应光信号 Involved in the light responsiveness
GT1-motif	GGTTAA	1	响应光信号 Involved in the light responsiveness
GT1-motif	GGTTAAT	1	响应光信号 Involved in the light responsiveness
MBS	CAACTG	1	响应干旱信号 Involved in the drought responsiveness
MYB	TAACCA	4	响应干旱信号 Involved in the drought responsiveness
TGACG-motif	TGACG	1	响应茉莉酸甲酯信号 Involved in the MeJA-responsiveness

Fig. 2 Expression characteristics of CaPUB18 gene in various tissues（A）and under heat stress（B）in pepper Different lowercase letters mean the significant difference at P < 0.05 level（n = 3）. The same below

Fig. 3 Subcellular localization of CaPUB18 A：Subcellular localization of CaPUB18；B：Colocalization of CaPUB18 with PIP2A，the marker protein of plasma membrane，under normal and high temperature respectively；C：Quantification of colocalization of CaPUB18with PIP2A under high temperatures

Fig. 4 Generation of CaPUB18-silenced pepper plants A：Phenotype of gene-silenced pepper plants；B：Test of silencing efficiency of CaPUB18，*** P < 0.001. TRV2:CaPUB18：CaPUB18-silenced plants；TRV2:CaPDS：Positive control；TRV2:00：Negative control. The same below

Fig. 5 Effect of CaPUB18-silenced on heat tolerance of pepper A：Phenotypes of pepper plants before and after heat-stress treatment；B-E：Relative electrical conductivity，malondialdehyde content，H2O2 accumulation and dead cells accumulation in pepper leaves before and after heat-stress treatment；F-G：Content of total chlorophyll in pepper detached leaf-discs before and after heat-stress treatment；H，I：Relative expression of heat-tolerance marker genes CaHSP70.1 and CaHsfA2 in pepper leaves before and after heat-stress treatment

Fig. 6 Identification and seedling survival rate of Arabidopsis thaliana lines CaPUB18-OE A and B：Identification of Arabidopsis CaPUB18-OE lines with semi-RT-PCR and qRT-PCR，respectively；C and D：Phenotypes and survival rate of 5-day-old Arabidopsis seedlings after heat-stress treatment with 45 ℃ for 50 min and then recovery with 22 ℃ for 3 d

Fig. 7 Phenotypes，relative electrical conductivity and MDA of Arabidopsis thaliana lines CaPUB18-OE A：Phenotypes of 21-day-old Arabidopsis plants after heat-stress treatment；B and C：Relative electrical conductivity，malondialdehyde content in Arabidopsis leaves before and after heat-stress treatment； Col-0：Arabidopsis wild type；OE-2，and OE-3：Arabidopsis CaPUB18-overexpression lines in Col-0

Fig. 8 Effects of CaPUB18-overexpression on heat tolerance of Arabidopsis A and D：Content of total chlorophyll in Arabidopsis detached leaf-discs before and after heat-stress treatment；B and E：Accumulation of H2O2（DAB staining）and mean gray values of 21-day-old Arabidopsis plants after heat-stress treatment；C and F：Dead cells（Trypan blue staining）and mean gray values of 21-day-old Arabidopsis plants after heat-stress treatment. Col-0：Arabidopsis wild type；OE-1，OE-2，and OE-3：Arabidopsis CaPUB18-overexpression lines in Col-0

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Functional Analysis of Pepper U-Box Domain-Containing Protein Gene CaPUB18 in Heat Tolerance

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