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

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

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

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

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 Online:2025-10-25 Published:2025-10-28
Contact: JING Danlong, GUO Qigao

Abstract

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

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.

Figures/Tables 11

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

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

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

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

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

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

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

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

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

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）

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

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