枇杷bHLH基因家族特征和EjbHLH104调控三倍体枇杷花期的功能分析

doi:10.16420/j.issn.0513-353x.2024-0790

园艺学报 ›› 2025, Vol. 52 ›› Issue (10): 2582-2596.doi: 10.16420/j.issn.0513-353x.2024-0790

• 遗传育种·种质资源·分子生物学 • 上一篇下一篇

枇杷bHLH基因家族特征和EjbHLH104调控三倍体枇杷花期的功能分析

夏燕¹, 胡若倩¹^,², 易广权¹, 孙洪文¹, 黄树欣¹, 池卓恒¹, 石敏³, 梁国鲁¹, 景丹龙¹^,^*(), 郭启高¹^,^*()

¹ 西南大学园艺园林学院，西南大学农业科学研究院，长江上游农业生物安全与绿色生产教育部重点实验室，三峡库区森林生态修复与利用重庆市重点实验室，重庆 400715
² 浙江大学农学与生物技术学院，农业部园艺植物生长发育与品质调控重点开放实验室，杭州 310058
³ 马边彝族自治县农业农村局，四川乐山 614600

收稿日期:2025-06-20 修回日期:2025-07-29 出版日期:2025-10-25 发布日期:2025-10-28
通讯作者:
^* E-mail：jingdanlong@swu.edu.cn，
qgguo@126.com
基金资助:
国家自然科学基金面上项目(32470386); 重庆市科学技术局项目(cstc2021jscx-gksbX0010); 重庆市科学技术局项目(CSTB2023TIAD-KPX0044); 重庆市科学技术局项目(CSTB2022NSCQ-MSX1541); 中央高校基本科研业务费能力提升项目(SWU-KT22055)

Characterization of Loquat bHLH Gene Family and Functional Analysis of EjbHLH104 on Flowering Regulation in Triploid Loquat

XIA Yan¹, HU Ruoqian¹^,², YI Guangquan¹, SUN Hongwen¹, HUANG Shuxin¹, CHI Zhuoheng¹, SHI Min³, LIANG Guolu¹, JING Danlong¹^,^*(), GUO Qigao¹^,^*()

¹ Chongqing Key Laboratory of Forest Ecological Restoration and Utilization in the Three Gorges Reservoir Area，Key Laboratory of Agricultural Biosafety and Green Production of Upper Yangtze River（Ministry of Education），Academy of Agricultural Sciences of Southwest University，College of Horticulture and Landscape Architecture，Southwest University，Chongqing 400715，China
² The State Agriculture Ministry Laboratory of Horticultural Plant Growth，Development and Quality Improvement，College of Agriculture & Biotechnology，Zhejiang University，Hangzhou 310058，China
³ Agricultural and Rural Bureau of Mabian Yi Autonomous County，Leshan，Sichuan 614600，China

Received:2025-06-20 Revised:2025-07-29 Published:2025-10-25 Online:2025-10-28

摘要/Abstract

摘要：

从枇杷（Eriobotrya japonica）基因组中鉴定到169个EjbHLH基因家族成员，这些基因在染色体上呈现不均等分布，划分为22个亚家族。采用RT-PCR技术克隆获得1个在早花三倍体枇杷‘无核早玉’花芽分化期显著高表达的bHLH成员EjbHLH104，其编码区序列长度为675 bp，编码224个氨基酸。蛋白质序列比对和分子系统发生分析表明，EjbHLH104具有典型且保守的bHLH保守结构域，与苹果MdbHLH104和白梨PbbHLH104等蔷薇科植物bHLH蛋白质的相似性较高，聚在真蔷薇分支Ⅰ。亚细胞定位分析表明，EjbHLH104蛋白质定位在细胞核，具有典型的bHLH转录因子亚细胞定位特性。实时荧光定量PCR分析表明，EjbHLH104基因在三倍体枇杷早花品种的花芽生理分化期和小花分化期表达量较高，在盛花期最高；而在二倍体早花品种的小花分化期表达量最高。EjbHLH104转基因拟南芥植株开花时间比野生型拟南芥显著提前，表明EjbHLH104基因过量表达具有促进拟南芥提前开花的功能。

关键词: 枇杷, 三倍体, bHLH基因家族, 功能分析, 花期调控

Abstract:

A total of 169 EjbHLH gene family members were identified from loquat genome，and were classified into 22 subfamilies. Then，the cDNA sequence of EjbHLH104，a member of the bHLH gene family，was isolated using RT-PCR from the flower buds of triploid loquat，and it was significantly increased in flower buds of early flowering triploid loquat‘Wuhe Zaoyu’. The coding sequence of EjbHLH104 was 675 bp，encoding 224 amino acids. Protein sequence alignment and phylogenetic tree analysis showed that EjbHLH104 has the conserved bHLH domain and is highly similar to MdbHLH104 of Malus × domestica and PbbHLH104 of Pyrus bretschneideri，which are clustered into the same branch of eurosidsⅠbHLH104 lineage. Subcellular localization analysis showed that the localization of EjbHLH104 protein was in the nucleus，suggesting it has subcellular localization characteristics of bHLH transcription factor. In early and late flowering cultivars of diploid and triploid loquats，the qRT-PCR analysis showed that expression levels of EjbHLH104 were higher at the stages of flower bud differentiation and floret differentiation and reached the highest at full bloom stage in the early flowering cultivars of triploid loquats. However，EjbHLH104 expression level was reached the highest at the development stage of floret differentiation in the early flowering cultivars of diploid loquat. Compared with the wild Arabidopsis thaliana，flowering time observation of EjbHLH104 transgenic Arabidopsis showed that overexpression of EjbHLH104 gene significantly exhibited early flowering，suggesting EjbHLH104 could promote flowering.

Key words: loquat, triploid, bHLH gene family, functional analysis, flowering regulation

夏燕, 胡若倩, 易广权, 孙洪文, 黄树欣, 池卓恒, 石敏, 梁国鲁, 景丹龙, 郭启高. 枇杷bHLH基因家族特征和EjbHLH104调控三倍体枇杷花期的功能分析[J]. 园艺学报, 2025, 52(10): 2582-2596.

XIA Yan, HU Ruoqian, YI Guangquan, SUN Hongwen, HUANG Shuxin, CHI Zhuoheng, SHI Min, LIANG Guolu, JING Danlong, GUO Qigao. Characterization of Loquat bHLH Gene Family and Functional Analysis of EjbHLH104 on Flowering Regulation in Triploid Loquat[J]. Acta Horticulturae Sinica, 2025, 52(10): 2582-2596.

图/表 11

图1 三倍体早花品种‘无核早玉’花芽发育阶段的形态 S1：花芽生理分化期；S2：花芽形态分化期；S3：花序主轴分化期；S4：花序支轴分化期；S5：花序侧生支轴的花芽快速伸长期；S6：小花分化期；S7：花蕾露白期；S8：盛花期。下同

Fig. 1 Morphological characterization of early-flowering triploid loquat‘Wuhe Zaoyu’ S1：Physiological differentiation of flower buds；S2：Morphological differentiation of flower buds；S3：Differentiation of the main inflorescence axis；S4：Differentiation of lateral inflorescence branches；S5：Rapid elongation of flower buds from lateral inflorescence branches；S6：Differentiation of florets；S7：White corollas of floral buds；S8：Full bloom. The same below

表1 4 个枇杷品种的花芽采样时间范围（2019—2021年）

Table 1 The acquisition time of the flower panicles of four loquat cultivars

时期 Stage	森尾早生 Moriowase	常白1号 Changbai 1	无核早玉 Wuhe Zaoyu	华玉无核1号 Huayu Wuhe 1
花芽生理分化期 Physiological differentiation of flower buds（S1）	07-04—07-12	07-06—07-15	07-04—07-12	07-06—07-15
花芽形态分化期 Morphological differentiation of flower buds（S2）	07-25—08-03	09-01—09-09	07-20—07-29	08-30—09-09
花序主轴分化期 Differentiation of the main inflorescence axis（S3）	08-21—08-30	09-22—09-30	08-18—08-25	09-21—09-30
花序支轴分化期 Differentiation of lateral inflorescence branches（S4）	09-04—09-12	10-12—10-15	08-29—09-07	10-10—10-15
花序侧生支轴的花芽快速伸长期 Rapid elongation of flower buds from lateral inflorescence branches（S5）	09-17—09-26	10-18—10-21	09-11—09-19	10-19—10-22
小花分化期 Differentiation of florets（S6）	10-04—10-09	10-25—10-29	09-22—09-29	10-25—10-30
花蕾露白期 White corollas of floral buds（S7）	10-13—10-21	11-02—11-08	10-05—10-09	11-04—11-11
盛花期 Full bloom（S8）	10-25—10-30	11-24—11-29	10-13—10-23	11-23—11-29

表2 EjbHLH104基因所用引物序列

Table 2 Primer sequences used for EjbHLH104 gene

引物名称 Primer name	序列（5′-3′） Sequence	用途 Purpose
EjbHLH104F	ATGGGGGAATGGATAGAGTAT	EjbHLH104 CDS全长扩增Amplification of EjbHLH104 CDS
EjbHLH104R	CTAAGCAGCGGGGGGCCTAA	EjbHLH104 CDS全长扩增Amplification of EjbHLH104 CDS
qEjbHLH104F	AGAGTGTCCCCGCAAGAG	实时荧光定量PCR Quantitative real-time PCR
qEjbHLH104R	GCATCATCAAGTATAGCAGCCT	实时荧光定量PCR Quantitative real-time PCR
qEjactinF	AATGGAACTGGAATGGTCAAGGC	定量PCR内参引物Primers for internal control
qEjactinR	TGCCAGATCTTCTCCATGTCATCCCA	定量PCR内参引物Primers for internal control
LEjbHLH104F	ggggtaccATGGGGGAATGGATAGAGTAT	亚细胞定位载体构建Vector construction of subcellular localization
LEjbHLH104R	gctctagaAGCAGCGGGGGGCCTAA	亚细胞定位载体构建Vector construction of subcellular localization
TEjbHLH104F	ggcgcgccATGGGGGAATGGATAGAGTAT	转基因拟南芥载体构建Vector construction of transgenic Arabidopsis
TEjbHLH104R	gctctagaCTAAGCAGCGGGGGGCCTAA	转基因拟南芥载体构建Vector construction of transgenic Arabidopsis
pFGC5941-35SF	TGAGACTTTTCAACAAAGGATAATT	转基因植株鉴定Identification of transgenic Arabidopsis
pFGC5941-35SR	TGTCCTCTCCAAATGAAATGAAC	转基因植株鉴定Identification of transgenic Arabidopsis

图2 bHLH基因家族成员在枇杷基因组上的分布

Fig. 2 Distribution of bHLH gene family members in loquat genome

图3 枇杷（Ej）和苹果（Md）bHLH家族成员的系统进化分析外圈绿色、黄色和罗马数字代表不同bHLH亚组。枇杷和苹果bHLH共聚类为24个亚家族。其中枇杷EjbHLH聚类在22个亚家族；图中红色箭头所示为EjbHLH104

Fig. 3 Phylogenetic tree analysis of loquat（Ej）and apple（Md）bHLH proteins Outer branches of green，yellow and Roman numbers are shown the different bHLH subgroups. The bHLH proteins of loquat and apple are classified into 24 subfamilies. EjbHLH are distributed among 22 subgroups. Red arrow indicates the EjbHLH104

图4 枇杷EjbHLH104基因克隆和编码蛋白结构特性分析 A：EjbHLH104基因在二、三倍体枇杷花芽分化期中的表达量（FPKM值），其中包括：晚花二倍体品种‘常白1号’，早花三倍体品种‘无核早玉’，晚花三倍体品种‘华玉无核1号’；B：三倍体枇杷‘无核早玉’花芽EjbHLH104的PCR扩增产物；C：EjbHLH104蛋白质二级结构，其中紫色表示不规则卷曲，蓝色表示α螺旋，红色表示延伸链，绿色表示β转角；D：EjbHLH104蛋白质三级结构线状结构模型；E：EjbHLH104蛋白质三级结构空间立体球模型

Fig. 4 Gene cloning of EjbHLH104 and structural characteristics of loquat EjbHLH104 protein A：FPKM values of flower bud differentiation in diploid and triploid cultivars，Changbai 1，late-flowering cultivar of diploid loquat，Wuhe Zaoyu，early-flowering cultivar of triploid loquats，Huayu Wuhe 1，late-flowering cultivar of triploid loquats；B：PCR amplification product of the EjbHLH104 gene from triploid loquat flower buds；C：Protein secondary structure of EjbHLH104，purple indicates random curls，blue indicates α helix，red indicates extended strand，green indicates β turn；D：Linear structure model of EjbHLH104 protein tertiary structure. E：Three-dimensional spatial structures of EjbHLH104

图5 枇杷EjbHLH104与其他植物bHLH104的系统进化分析

Fig. 5 Phylogenetic analysis of EjbHLH104 and related members of other plants bHLH104

图6 枇杷EjbHLH104与其他植物bHLH104同源蛋白质序列比较 bHLH104同源蛋白质序列结构域划分参考Carretero-Paulet 等（2010）和Wu等（2022）。basic：碱性结构域；Helix 1：螺旋结构域1；Loop：环形结构域；Helix 2：螺旋结构域2。Ej：枇杷，Md：苹果，Pb：白梨，Ms：森林苹果，Pa：欧洲甜樱桃，Pm：梅花，Fv：森林草莓，Rr：玫瑰，Rc：月季

Fig. 6 Comparison of amino acid sequences between loquat EjbHLH104 and related members of other plants bHLH104 The underlined regions of EjbHLH104 were according to Carretero-Paulet et al.（2010）and Wu et al.（2022). basic：Basic domain；Helix 1：Helix 1 domain；Loop：Loop domain；Helix 2：Helix 2 domain. Ej：Eriobotrya japonica，Md：Malus × domestica，Pb：Pyrus bretschneideri，Ms：Malus sylvestris，Pa：Prunus avium，Pm：Prunus mume，Fv：Fragaria vesca，Rr：Rosa rugosa，Rc：Rosa chinensis

图7 枇杷35S::EjbHLH104-GFP基因在烟草细胞中的亚细胞定位

Fig. 7 Subcellular localization of 35S::EjbHLH104-GFP in tobacco cell

图8 EjbHLH104基因在二倍体和三倍体早花（A、C）晚花（B、D）枇杷中的相对表达量不同小写字母表示表达量差异显著（P < 0.05）

Fig. 8 Relative expression levels of EjbHLH104 in early（A，C）and late（B，D）flowering varieties on diploid and triploid loquat Significant differences are indicated by different lowercase letters（P < 0.05）

图9 EjbHLH104转基因拟南芥开花表型分析 A：除草剂筛选；B：阳性植株的PCR鉴定（M为DNA marker，WT为野生型拟南芥，1 ~ 23为筛选的阳性植株，P为含目的基因超表达载体质粒）；C：开花表型；D：开花时间比较。**表示开花时间差异极显著（P < 0.01）

Fig. 9 Observation of flowering time phenotypes in EjbHLH104 transgenic Arabidopsis A：Screening of transgenic plants using herbicide；B：PCR verification of transgenic Arabidopsis（M indicates DNA marker，WT indicates wild-type Arabidopsis，1-23 indicates transgenic Arabidopsis lines，P indicates the PCR product with plasmid containing EjbHLH104 as a template）；C：Flowering phenotypes；D：Flowering time analysis.** indicates significant difference in days of flowering at P < 0.01

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枇杷bHLH基因家族特征和EjbHLH104调控三倍体枇杷花期的功能分析

Characterization of Loquat bHLH Gene Family and Functional Analysis of EjbHLH104 on Flowering Regulation in Triploid Loquat

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枇杷bHLH基因家族特征和EjbHLH104调控三倍体枇杷花期的功能分析

Characterization of Loquat bHLH Gene Family and Functional Analysis of EjbHLH104 on Flowering Regulation in Triploid Loquat

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相关文章 15

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