草莓HSP20基因家族的鉴定及在果实高温胁迫响应与成熟调控中的功能分析

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

园艺学报 ›› 2026, Vol. 53 ›› Issue (7): 2053-2074.doi: 10.16420/j.issn.0513-353x.2026-0341

• 遗传育种 · 种质资源 · 分子生物学 • 上一篇下一篇

草莓HSP20基因家族的鉴定及在果实高温胁迫响应与成熟调控中的功能分析

赵艳侠¹, 孙家波¹, 吴依晴¹^,², 李倩², 梁红敏¹, 李冰冰²^,^*()

¹ 山东省农业科学院休闲农业研究所，农业农村部华东都市农业重点实验室，生态园艺植物育种山东省工程研究中心，济南 250100
² 中国农业大学园艺学院，北京 100193

收稿日期:2026-04-13 修回日期:2026-06-22 出版日期:2026-07-25 发布日期:2026-07-23
通讯作者:
^* E-mail：libingbing@cau.edu.cn
基金资助:
山东省农业科学院农业科技创新工程项目(CXGC2026B30); 山东省农业科学院农业科技创新工程项目(CXGC2026H22); 新疆维吾尔自治区科技援疆项目(2026E02025); 国家重点研发计划项目(2023YFF1001700); 国家自然科学基金面上项目(32572990); 中国农业大学2115人才培育发展支持计划和中国乡村发展基金会新长城科技小院助力计划（山东历城草莓科技小院）项目(10122424005)

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 Published:2026-07-25 Online:2026-07-23

摘要/Abstract

摘要：

小分子热激蛋白20（HSP20）在植物高温逆境应答与生长发育调控中发挥关键作用。为解析草莓HSP20基因家族的高温响应机制及其对果实成熟的调控功能，本研究中以二倍体森林草莓（Fragaria vesca）为材料，开展了FvHSP20基因家族全基因组鉴定，系统分析其基因结构、系统发育、染色体定位与顺式作用元件，并结合转录组与qRT-PCR技术解析该基因家族在森林草莓和栽培草莓（Fragaria × ananassa）不同果实发育阶段及高温胁迫下的表达模式，同时在栽培草莓中验证关键基因的功能。结果表明：在森林草莓基因组中共鉴定出33个FvHSP20家族成员，其编码蛋白氨基酸数量为78 ~ 399，编码蛋白的理化性质稳定，亚细胞定位于细胞质、叶绿体、线粒体等细胞器，存在明显区室化特征。系统进化分析结果显示，家族基因可划分为13个亚族，并新命名其中一支为CXⅡ亚族；草莓中缺失内质网（ER）亚族成员，呈现物种进化特异性。33个基因不均匀分布于7条染色体，多数基因结构简单，仅含0 ~ 1个内含子。各基因启动子均含胁迫与激素响应元件，以光响应、脱落酸与茉莉酸甲酯调控相关元件最为丰富。FvHSP20/FaHSP20表达具有果实发育阶段特异性，白果期对高温胁迫敏感，HSP20-13/14/20等基因在高温处理后显著上调；不同栽培品种的响应模式存在差异，耐热品种‘香野’转录表达更平稳。瞬时过表达试验证实，FaHSP20-13在常温条件下能促进草莓果实提前着色、增强糖分积累；高温环境下该基因维持脱落酸信号稳态，缓解高温对果实发育的抑制，兼具促成熟与耐热的功能。本研究中明确了草莓HSP20家族的分子特征与高温应答特点，挖掘出关键耐热与成熟调控候选基因，可为草莓耐高温分子育种和品质改良提供理论依据与基因资源。

关键词: 草莓, HSP20, 全基因组鉴定, 高温胁迫, 果实成熟调控

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

赵艳侠, 孙家波, 吴依晴, 李倩, 梁红敏, 李冰冰. 草莓HSP20基因家族的鉴定及在果实高温胁迫响应与成熟调控中的功能分析[J]. 园艺学报, 2026, 53(7): 2053-2074.

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.

图/表 15

表1 qRT-PCR引物序列

Table 1 qRT-PCR primer sequences

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

表2 草莓基因组中FvHSP20基因家族成员理化性质

Table 2 Physicochemical of characteristics of FvHSP20 gene family members in the strawberry genome

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

表2 草莓基因组中FvHSP20基因家族成员理化性质

Table 2 Physicochemical of characteristics of FvHSP20 gene family members in the strawberry genome

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

图1 拟南芥（At）、水稻（Os）和草莓（Fv）中HSP20蛋白的系统发育关系

Fig. 1 Phylogenetic relationship of HSP20 proteins in Arabidopsis thaliana（At），Oryza sativa（Os），and Fragaria vesca（Fv）

图2 森林草莓FvHSP20基因家族成员在染色体上的分布

Fig. 2 Chromosomal distribution of FvHSP20 gene family members of Fragaria vesca

图3 FvHSP20的系统发育关系（A）、基因结构（B）、保守蛋白基序（C）及保守结构域（D）

Fig. 3 Phylogenetic relationships（A），gene structures（B），conserved protein motifs（C） and conserved domain（D） of the FvHSP20

表3 FvHSP20保守的基序序列

Table 3 Conserved motif sequences of FvHSP20

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

图4 FvHSP20基因启动子中预测的顺式调控元件 A：顺式调控元件的位置和种类；B：顺式调控元件数量统计

Fig. 4 Predicted cis-elements in FvHSP20 promoters A：Location and types of cis-regulatory elements；B：Statistical count of cis-regulatory elements

图5 FvHSP20基因在不同果实发育阶段的表达模式分析不同小写字母表示处理间差异显著（P < 0.05，n = 3）。下同

Fig. 5 Analysis of FvHSP20 gene expression patterns at different fruit developmental stages Different lowercase letters indicate significant differences among different treatments（P < 0.05，n = 3）. The same as below

图6 高温时FvHSP20基因在白果期（W）和转色期（T）的表达模式分析 CK：25 ℃，对照；HT：35 ℃高温胁迫。下同

Fig. 6 Analysis of FvHSP20 gene expression patterns during the white fruit stage（W）and turning color stage（T）under high-temperature CK：25 ℃，control；HT：35 ℃，high-temperature. The same below

图7 ‘红颜’和‘香野’白果期草莓果实在35 ℃高温条件下表型变化

Fig. 7 Phenotypic changes of‘Benihoppe’ and‘Kaorino’ strawberry fruits at the white fruit stage under 35 ℃ high temperature

图8 草莓‘红颜’（HY）和‘香野’（XY）白果期FaHSP20基因在高温（35 ℃）下的表达模式分析

Fig. 8 Analysis of FvHSP20 gene expression patterns in the white stage of‘Benihoppe’（HY）and‘Kaorino’（XY）strawberrys under high-temperature（35 ℃）

图9 高温胁迫下草莓果实不同发育阶段FaHSP20关键基因的表达模式分析

Fig. 9 Analysis of the expression patterns of key FaHSP20 genes in strawberry fruits at different developmental stages under high-temperature stress

图10 田间果实中FaHSP20过表达的功能分析 A：果实成熟进程变化；B：不同发育时期果实比例；C：3个关键基因过表达后目的基因表达水平；D ~ F：侵染12 d后的果实花色苷、原花青素含量以及硬度

Fig. 10 Functional analysis of overexpression of FaHSP20 gene in fruits in the field A：Changes in fruit ripening process；B：Statistical percentage of fruit developmental stages；C：Expression levels of target genes after overexpression of three FaHSP20 genes；D-F：Anthocyanin，proanthocyanidin contents and firmness of fruits after 12 days of infection

图11 常温下离体果实FaHSP20-13基因过表达的功能分析 A：果实成熟进程；B ~ D：离体侵染5 d后果实中的花色苷、可溶性糖以及脱落酸醛（ABA-ald）、脱落酸（ABA）、脱落酸葡萄糖酯（ABA-GE）含量；E：离体侵染5 d后目的基因和成熟相关基因的表达

Fig. 11 Functional analysis of FaHSP20-13 gene overexpression at room temperature A：Changes in the ripening process；B-D：Contents of anthocyanins，soluble sugars，abscisic aldehyde（ABA-ald），abscisic acid（ABA）and abscisic acid glucose ester（ABA-GE）after 5 days of in vitro inoculation；E：Expression levels of target genes and ripening-related genes in fruits after 5 days of in vitro inoculation

图12 高温胁迫下离体果实FaHSP20-13基因过表达的功能分析 A：果实成熟进程；B ~ D：侵染5 d后果实花青素、糖以及ABA-ald、ABA、ABA-GE含量；E：离体侵染5 d后目的基因和成熟相关基因的变化

Fig. 12 Functional analysis of transient overexpression of FaHSP20-13 gene under high-temperature stress A：Changes in the ripening process；B-D：Contents of anthocyanins，soluble sugars，ABA-ald，ABA and ABA-GE in fruits after 5 days of in vitro inoculation；E：Expression levels of target genes and ripening-related genes in fruits after 5 days of in vitro inoculation

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草莓HSP20基因家族的鉴定及在果实高温胁迫响应与成熟调控中的功能分析

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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草莓HSP20基因家族的鉴定及在果实高温胁迫响应与成熟调控中的功能分析

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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