Genome-Wide Identification of the HSP20 Gene Family in Strawberry and its Functional Analysis in Fruit Response to High-Temperature Stress and Ripening Regulation

doi:10.16420/j.issn.0513-353x.2026-0341

Acta Horticulturae Sinica ›› 2026, Vol. 53 ›› Issue (7): 2053-2074.doi: 10.16420/j.issn.0513-353x.2026-0341

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

Genome-Wide Identification of the HSP20 Gene Family in Strawberry and its Functional Analysis in Fruit Response to High-Temperature Stress and Ripening Regulation

ZHAO Yanxia¹, SUN Jiabo¹, Wu Yiqing¹^,², LI Qian², LIANG Hongmin¹, LI Bingbing²^,^*()

¹ Institute of Leisure Agriculture， Shandong Academy of Agricultural Sciences;Key Laboratory of East China Urban Agriculture，Ministry of Agriculture and Rural Affairs;Shandong Engineering Research Center of Ecological Horticultural Plant Breeding， Ji’nan 250100， China
² College of Horticulture， China Agricultural University， Beijing 100193， China

Received:2026-04-13 Revised:2026-06-22 Online:2026-07-25 Published:2026-07-23
Contact: LI Bingbing

Abstract

Abstract:

Small heat shock protein 20（HSP20）are pivotal regulators involved in plant high-temperature stress response as well as growth and development. To dissect the high-temperature response mechanism of the strawberry HSP20 gene family and its regulatory function in fruit ripening，diploid woodland strawberry（Fragaria vesca）was used as material in this study. A genome-wide identification of the FvHSP20 gene family was carried out using bioinformatics methods，and the gene structure，phylogeny，chromosomal localization，and cis-acting elements were systematically analyzed. Furthermore，combined with transcriptome and qRT-PCR technologies，the expression patterns of this family at different fruit developmental stages and under high-temperature stress in both woodland strawberry and cultivated strawberry（Fragaria × ananassa）were resolved，and the biological function of key gene was further validated in cultivated strawberry. The results showed that a total of 33 FvHSP20 genes were identified from the F. vesca genome. The encoded HSP20 proteins contained 78 to 399 amino acids with stable physicochemical properties，and were targeted to cytoplasm，chloroplasts，mitochondria and other organelles，exhibiting obvious subcellular compartmentalization. Phylogenetic analysis categorized these genes into 13 subfamilies，among which a novel clade designated CXⅡ subfamily was newly classified in this study. Notably，strawberry HSP20 family lacked endoplasmic reticulum subfamily members，reflecting unique evolutionary characteristics of this species. The 33 FvHSP20 genes were unevenly mapped onto seven strawberry chromosomes，most family members possessed compact gene structures with zero or one intron. Promoter sequence of various genes analysis revealed numerous stress-responsive and hormone-responsive cis-elements，among which light-responsive，abscisic acid（ABA）-responsive and methyl jasmonate（MeJA）-responsive elements were the most abundant. Transcriptomic results indicated that both FvHSP20 and FaHSP20 genes displayed fruit developmental stage-specific expression patterns. Fruits at the white stage were most susceptible to high-temperature stress，and core genes including HSP20-13，HSP20-14 and HSP20-20 were significantly upregulated upon heat treatment. Distinct transcriptional responses to heat stress were detected between two cultivated strawberry cultivars：the heat-tolerant cultivar‘Kaorino’exhibited milder and more stable gene expression fluctuations. Transient overexpression assays verified that FaHSP20-13 accelerated fruit coloration，enhanced sugar enrichment under normal temperature conditions. Furthermore，FaHSP20-13 maintained ABA signaling homeostasis to relieve heat-induced growth inhibition in strawberry fruits，performing dual functions in promoting maturation and heat resistance. In this study，the basic characteristics and high-temperature response patterns of the strawberry HSP20 family were clarified，and key candidate genes for heat tolerance and ripening regulation were identified，which provided theoretical basis and genetic resources for molecular breeding of heat tolerance and high-quality improvement in strawberry.

Key words: strawberry, HSP20, genome-wide identification, high-temperature stress, fruit ripening regulation

ZHAO Yanxia, SUN Jiabo, Wu Yiqing, LI Qian, LIANG Hongmin, LI Bingbing. Genome-Wide Identification of the HSP20 Gene Family in Strawberry and its Functional Analysis in Fruit Response to High-Temperature Stress and Ripening Regulation[J]. Acta Horticulturae Sinica, 2026, 53(7): 2053-2074.

Figures/Tables 15

References 57

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基因 Gene	上游引物序列（5′-3′） Forward primer	下游引物序列（5′-3′） Reverse primer
FaSUS1	CCCTGATTCTGACCTTTACTGG	GATGATGAAGTCGGTGTGGTT
FaSUS2	TTGCTGAGGATGTTGCAGGT	AACAACGAGGCGACGAGATT
FaSPS1	GAGCCTTGAATGTCCCAATGT	ATCTCCTGTCTGGTGCTGGTT
FaSPS3	CGTAGATTGGAGTTATGGAGAGC	CGAATGATGTAAGAACCACTGC
FaCHS1	GCCTTTGTTTGGCTGGCT	CCCAGGAACATCTTTGAGGA
FaCHI1	AGCGAAAGCCATTGAAAAGT	CATTTGGTGATTGTGTGAAGAG
FaPAL	CTTCTGTGGTGCTGTTTGATG	AGGGTGGTGCTTCAGTTTATGT
FaMYB10	CAACAGCACCACCACAGACT	GCTTGCCGATTGTACCGTAT
FaPYL1	GCTTCGTAATGAGCGTGGGA	CTCTCCTCGTCCAGCAAGTC
FaCYP707A4a	CCCGCTACAAAGAAGAACTGA	TGTCACCGATAATGTGCCTC
FaNCED1	ACTGCTTCTGCTTCCATCTCT	AGACACTCGTCGCATTCATT
FaNCED2	GACACCTTTTGCTTCCACTTG	GGATTTCGGACAACACACTCT
FaNCED3	CAAACCCTATCACCCTTCTCAC	TGGGCAACTTCTGCTTCTTT
FaAAO3	AACAGCTTCTTCAATGGGGAGG	TCAGTTTCTACCTGCGAACCA
FaRIF	GAGCACCGGAAATGGTTCTA	CCCCGGAAGTTGATACTGTG
FaHSP20-13	GGACCCATTCCTCGACTTCC	GCTGCTTTGAGAACGTGAGC
FaHSP20-14	TGTCGCTCATCCCCAGTTTC	GACAAAGGCCGAGTTTTCGC
FaHSP20-20	CAATCGCTCTCAGAAGGGCA	CGAGAAACAGAGGGGGCTAC
FaACTIN	GCCAACCGTGAGAAGATG	TCCAGAGTCAAGAACAATACCAG

基因 Gene	上游引物序列（5′-3′） Forward primer	下游引物序列（5′-3′） Reverse primer
FaSUS1	CCCTGATTCTGACCTTTACTGG	GATGATGAAGTCGGTGTGGTT
FaSUS2	TTGCTGAGGATGTTGCAGGT	AACAACGAGGCGACGAGATT
FaSPS1	GAGCCTTGAATGTCCCAATGT	ATCTCCTGTCTGGTGCTGGTT
FaSPS3	CGTAGATTGGAGTTATGGAGAGC	CGAATGATGTAAGAACCACTGC
FaCHS1	GCCTTTGTTTGGCTGGCT	CCCAGGAACATCTTTGAGGA
FaCHI1	AGCGAAAGCCATTGAAAAGT	CATTTGGTGATTGTGTGAAGAG
FaPAL	CTTCTGTGGTGCTGTTTGATG	AGGGTGGTGCTTCAGTTTATGT
FaMYB10	CAACAGCACCACCACAGACT	GCTTGCCGATTGTACCGTAT
FaPYL1	GCTTCGTAATGAGCGTGGGA	CTCTCCTCGTCCAGCAAGTC
FaCYP707A4a	CCCGCTACAAAGAAGAACTGA	TGTCACCGATAATGTGCCTC
FaNCED1	ACTGCTTCTGCTTCCATCTCT	AGACACTCGTCGCATTCATT
FaNCED2	GACACCTTTTGCTTCCACTTG	GGATTTCGGACAACACACTCT
FaNCED3	CAAACCCTATCACCCTTCTCAC	TGGGCAACTTCTGCTTCTTT
FaAAO3	AACAGCTTCTTCAATGGGGAGG	TCAGTTTCTACCTGCGAACCA
FaRIF	GAGCACCGGAAATGGTTCTA	CCCCGGAAGTTGATACTGTG
FaHSP20-13	GGACCCATTCCTCGACTTCC	GCTGCTTTGAGAACGTGAGC
FaHSP20-14	TGTCGCTCATCCCCAGTTTC	GACAAAGGCCGAGTTTTCGC
FaHSP20-20	CAATCGCTCTCAGAAGGGCA	CGAGAAACAGAGGGGGCTAC
FaACTIN	GCCAACCGTGAGAAGATG	TCCAGAGTCAAGAACAATACCAG

基因名 Gene name	基因ID Gene ID	染色体 Chromo- some	氨基酸数量 No. of amino acids	分子量/kD Molecular weight	等电点 pI	稳定系数 Instability index	脂溶性指数Aliphatic index	亲水性 Hydro- phylicity	亚细胞定位 Subcellular localization
FvHSP20-1	FvH4_1g07080.t1	Chr1	142	16.02	6.92	58.51	83.73	-0.434	过氧化物酶体 Peroxisomal
FvHSP20-2	FvH4_1g11472.t1	Chr1	246	27.23	9.36	59.87	82.85	-0.550	叶绿体Chloroplast
FvHSP20-3	FvH4_2g05762.t1	Chr2	146	16.43	4.98	47.35	79.45	-0.566	细胞质Cytoplasmic
FvHSP20-4	FvH4_2g31370.t1	Chr2	240	26.30	9.13	48.26	74.21	-0.685	细胞质Cytoplasmic
FvHSP20-5	FvH4_2g31371.t1	Chr2	159	17.82	5.34	42.49	85.22	-0.443	细胞质Cytoplasmic
FvHSP20-6	FvH4_3g02330.t1	Chr3	141	16.52	5.36	61.28	87.80	-0.448	细胞质Cytoplasmic
FvHSP20-7	FvH4_3g13900.t1	Chr3	150	17.09	7.06	45.99	80.47	-0.702	细胞质Cytoplasmic
FvHSP20-8	FvH4_3g21680.t1	Chr3	196	22.62	5.38	48.97	69.59	-0.671	细胞质Cytoplasmic
FvHSP20-9	FvH4_3g27040.t2	Chr3	399	44.61	9.00	45.62	66.49	-0.768	细胞质Cytoplasmic
FvHSP20-10	FvH4_3g29131.t1	Chr3	90	10.25	9.98	52.94	75.78	-0.939	线粒体Mitochondrion
FvHSP20-11	FvH4_3g35270.t1	Chr3	151	17.31	6.19	51.03	78.68	-0.685	细胞核Nucleus
FvHSP20-12	FvH4_3g35280.t1	Chr3	107	12.34	9.39	55.35	65.61	-0.462	细胞质Cytoplasmic
FvHSP20-13	FvH4_3g37320.t1	Chr3	143	15.96	4.69	58.01	87.90	-0.215	细胞质Cytoplasmic
FvHSP20-14	FvH4_3g37330.t1	Chr3	157	18.13	5.35	53.60	73.18	-0.641	细胞核Nucleus
FvHSP20-15	FvH4_3g37360.t1	Chr3	154	17.81	5.51	38.91	70.19	-0.737	细胞核Nucleus
FvHSP20-16	FvH4_3g38620.t1	Chr3	158	18.09	5.81	48.23	72.09	-0.685	细胞核Nucleus
FvHSP20-17	FvH4_4g24850.t1	Chr4	232	26.22	9.24	57.46	62.24	-0.759	叶绿体Chloroplast
FvHSP20-18	FvH4_5g15440.t1	Chr5	247	27.59	5.18	36.12	81.26	-0.552	细胞质Cytoplasmic
FvHSP20-19	FvH4_5g18340.t1	Chr5	226	25.37	8.70	42.60	73.36	-0.700	细胞质Cytoplasmic
FvHSP20-20	FvH4_5g18720.t1	Chr5	225	25.12	7.82	67.05	77.07	-0.616	线粒体Mitochondrial
FvHSP20-21	FvH4_6g02120.t1	Chr6	207	22.88	6.85	55.74	75.85	-0.546	叶绿体Chloroplast
FvHSP20-22	FvH4_6g02150.t1	Chr6	101	11.51	8.78	48.72	66.53	-0.791	叶绿体Chloroplast
FvHSP20-23	FvH4_6g21590.t1	Chr6	170	18.71	5.14	50.74	73.35	-0.656	线粒体Mitochondrial
FvHSP20-24	FvH4_6g29880.t1	Chr6	229	25.53	5.90	49.19	77.51	-0.694	线粒体Mitochondrial
FvHSP20-25	FvH4_6g35450.t1	Chr6	149	16.47	7.78	41.89	88.26	-0.505	细胞核Nucleus
FvHSP20-26	FvH4_6g52210.t1	Chr6	141	15.96	5.78	40.45	84.18	-0.712	细胞核Nucleus
FvHSP20-27	FvH4_7g04830.t1	Chr7	170	19.00	8.82	58.50	78.59	-0.389	细胞质Cytoplasmic
FvHSP20-28	FvH4_7g05240.t1	Chr7	382	42.29	9.04	37.33	67.98	-0.839	细胞质Cytoplasmic
FvHSP20-29	FvH4_7g19240.t1	Chr7	78	8.95	9.77	68.28	94.87	-0.429	叶绿体Chloroplast
FvHSP20-30	FvH4_7g23650.t1	Chr7	158	17.94	9.10	48.89	73.35	-0.692	细胞质Cytoplasmic
FvHSP20-31	FvH4_7g23710.t3	Chr7	184	20.95	9.10	45.78	60.92	-0.860	细胞质Cytoplasmic
FvHSP20-32	FvH4_7g27540.t1	Chr7	199	22.41	6.01	61.37	79.75	-0.539	线粒体Mitochondrial
FvHSP20-33	FvH4_7g32800.t1	Chr7	227	27.07	6.64	55.24	71.32	-0.634	叶绿体Chloroplast

基因名 Gene name	基因ID Gene ID	染色体 Chromo- some	氨基酸数量 No. of amino acids	分子量/kD Molecular weight	等电点 pI	稳定系数 Instability index	脂溶性指数Aliphatic index	亲水性 Hydro- phylicity	亚细胞定位 Subcellular localization
FvHSP20-1	FvH4_1g07080.t1	Chr1	142	16.02	6.92	58.51	83.73	-0.434	过氧化物酶体 Peroxisomal
FvHSP20-2	FvH4_1g11472.t1	Chr1	246	27.23	9.36	59.87	82.85	-0.550	叶绿体Chloroplast
FvHSP20-3	FvH4_2g05762.t1	Chr2	146	16.43	4.98	47.35	79.45	-0.566	细胞质Cytoplasmic
FvHSP20-4	FvH4_2g31370.t1	Chr2	240	26.30	9.13	48.26	74.21	-0.685	细胞质Cytoplasmic
FvHSP20-5	FvH4_2g31371.t1	Chr2	159	17.82	5.34	42.49	85.22	-0.443	细胞质Cytoplasmic
FvHSP20-6	FvH4_3g02330.t1	Chr3	141	16.52	5.36	61.28	87.80	-0.448	细胞质Cytoplasmic
FvHSP20-7	FvH4_3g13900.t1	Chr3	150	17.09	7.06	45.99	80.47	-0.702	细胞质Cytoplasmic
FvHSP20-8	FvH4_3g21680.t1	Chr3	196	22.62	5.38	48.97	69.59	-0.671	细胞质Cytoplasmic
FvHSP20-9	FvH4_3g27040.t2	Chr3	399	44.61	9.00	45.62	66.49	-0.768	细胞质Cytoplasmic
FvHSP20-10	FvH4_3g29131.t1	Chr3	90	10.25	9.98	52.94	75.78	-0.939	线粒体Mitochondrion
FvHSP20-11	FvH4_3g35270.t1	Chr3	151	17.31	6.19	51.03	78.68	-0.685	细胞核Nucleus
FvHSP20-12	FvH4_3g35280.t1	Chr3	107	12.34	9.39	55.35	65.61	-0.462	细胞质Cytoplasmic
FvHSP20-13	FvH4_3g37320.t1	Chr3	143	15.96	4.69	58.01	87.90	-0.215	细胞质Cytoplasmic
FvHSP20-14	FvH4_3g37330.t1	Chr3	157	18.13	5.35	53.60	73.18	-0.641	细胞核Nucleus
FvHSP20-15	FvH4_3g37360.t1	Chr3	154	17.81	5.51	38.91	70.19	-0.737	细胞核Nucleus
FvHSP20-16	FvH4_3g38620.t1	Chr3	158	18.09	5.81	48.23	72.09	-0.685	细胞核Nucleus
FvHSP20-17	FvH4_4g24850.t1	Chr4	232	26.22	9.24	57.46	62.24	-0.759	叶绿体Chloroplast
FvHSP20-18	FvH4_5g15440.t1	Chr5	247	27.59	5.18	36.12	81.26	-0.552	细胞质Cytoplasmic
FvHSP20-19	FvH4_5g18340.t1	Chr5	226	25.37	8.70	42.60	73.36	-0.700	细胞质Cytoplasmic
FvHSP20-20	FvH4_5g18720.t1	Chr5	225	25.12	7.82	67.05	77.07	-0.616	线粒体Mitochondrial
FvHSP20-21	FvH4_6g02120.t1	Chr6	207	22.88	6.85	55.74	75.85	-0.546	叶绿体Chloroplast
FvHSP20-22	FvH4_6g02150.t1	Chr6	101	11.51	8.78	48.72	66.53	-0.791	叶绿体Chloroplast
FvHSP20-23	FvH4_6g21590.t1	Chr6	170	18.71	5.14	50.74	73.35	-0.656	线粒体Mitochondrial
FvHSP20-24	FvH4_6g29880.t1	Chr6	229	25.53	5.90	49.19	77.51	-0.694	线粒体Mitochondrial
FvHSP20-25	FvH4_6g35450.t1	Chr6	149	16.47	7.78	41.89	88.26	-0.505	细胞核Nucleus
FvHSP20-26	FvH4_6g52210.t1	Chr6	141	15.96	5.78	40.45	84.18	-0.712	细胞核Nucleus
FvHSP20-27	FvH4_7g04830.t1	Chr7	170	19.00	8.82	58.50	78.59	-0.389	细胞质Cytoplasmic
FvHSP20-28	FvH4_7g05240.t1	Chr7	382	42.29	9.04	37.33	67.98	-0.839	细胞质Cytoplasmic
FvHSP20-29	FvH4_7g19240.t1	Chr7	78	8.95	9.77	68.28	94.87	-0.429	叶绿体Chloroplast
FvHSP20-30	FvH4_7g23650.t1	Chr7	158	17.94	9.10	48.89	73.35	-0.692	细胞质Cytoplasmic
FvHSP20-31	FvH4_7g23710.t3	Chr7	184	20.95	9.10	45.78	60.92	-0.860	细胞质Cytoplasmic
FvHSP20-32	FvH4_7g27540.t1	Chr7	199	22.41	6.01	61.37	79.75	-0.539	线粒体Mitochondrial
FvHSP20-33	FvH4_7g32800.t1	Chr7	227	27.07	6.64	55.24	71.32	-0.634	叶绿体Chloroplast

基序Motif	基序一致性序列Motif consensus
1	FSRRFRLPENAKVDEIKAEMENGVLTVT
2	DWKETPEAHVFKADLPGLKKEDVKVEVED
3	GVLQISGERKREEEE
4	FSLDLWDPFKDFPFPSSSLST
5	KWHRVERSSGK
6	FPSPPRRLPALRAQAGGDGSVDVAVNQGSNNQGTQMEKRPSARR
7	PKVEVKKPDVKAIEI
8	RENSAFVNTRI
9	RTLSQMLDTMDRJFEBPMLSPG
10	MSLIPSFRRNS

Genome-Wide Identification of the HSP20 Gene Family in Strawberry and its Functional Analysis in Fruit Response to High-Temperature Stress and Ripening Regulation

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