Scientia Agricultura Sinica ›› 2026, Vol. 59 ›› Issue (8): 1748-1759.doi: 10.3864/j.issn.0578-1752.2026.08.011

• HORTICULTURE • Previous Articles     Next Articles

Physiological Mechanism and Relevant Gene Identification for Fruit Enlargement in Satsuma mandarin+Taoye sweet orange Somatic Cybrid

GUAN ShuPing1(), LI Ke1(), XIE KaiDong1, DUAN YaoYuan1, WANG TingTing1, ZHOU ZhiHu2, WU XiaoMeng1, GUO WenWu1,*()   

  1. 1 College of Horticulture & Forestry Sciences, Huazhong Agricultural University/National Key Laboratory for Germplasm Innovation & Utilization of Horticultural Crops, Wuhan 430070
    2 Agricultural and Rural Service Center of Qu Yuan Town, Zigui County, Zigui 443600, Hubei
  • Received:2025-11-19 Accepted:2026-01-14 Online:2026-04-16 Published:2026-04-21
  • Contact: GUO WenWu

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Abstract:

【Objective】Using the somatic cybrid Satsuma mandarin + Taoye sweet orange (G1+TYC), generated by protoplast fusion, and its mesophyll parent Taoye sweet orange (TYC) as materials, this study aimed to elucidate the physiological mechanism underlying fruit enlargement in G1+TYC and to identify candidate genes associated with the increased fruit size observed in this somatic cybrid.【Method】Ovaries and pulp tissues of G1+TYC and TYC at different developmental stages were used as materials. Dynamic monitoring of fruit size traits, determination of sugar and acid contents, and measurement of endogenous hormone levels were performed across various fruit developmental stages including the mature stage. Combined with transcriptome and bioinformatics analysis, this study was conducted to clarify the physiological causes of fruit enlargement in G1+TYC and to identify key candidate genes controlling fruit size.【Result】Dynamic monitoring of fruit development in three consecutive years showed that the transverse diameter of G1+TYC fruits was significantly larger than that of TYC starting from 75 days after flower (DAF), and the longitudinal diameter exhibited a significant superiority from 105 DAF onwards, with these differences persisting until 240 DAF. Taking the 2022 data as an example, the transverse and longitudinal diameters of the fruits from G1+TYC reached 79.14 and 66.75 mm at 240 DAF, respectively, while those from TYC were only 56.02 and 53.20 mm during the same period. This directly reflects the significant enlargement of fruit diameter of G1+TYC compared with TYC. The mature fruits of G1+TYC also had significantly higher fruit weight, segment width, single juice sac weight and area than TYC, indicating that the enlargement of juice sac volume contributed to the increase in segment width, thereby promoting the overall fruit size expansion of G1+TYC. Sugar and acid content determination revealed that the contents of fructose, glucose and sucrose in G1+TYC pulp were significantly higher than those in TYC during 90-240 DAF, whereas the contents of citric acid and total acids were significantly lower in G1+TYC. In terms of other organic acids, the difference in quinic acid and malic acid content between the two genotypes showed no obvious regularity. Endogenous hormone analysis demonstrated that the IAA content in G1+TYC pulp was significantly higher than that in TYC during 30-60 DAF. The GA3 content of G1+TYC was significantly higher only at 0 DAF, while the ZT content showed poor repeatability across the two years of measurement. The ABA content increased rapidly after 90 DAF, and the ABA content in G1+TYC pulp was significantly higher than that in TYC at 180 DAF. Combined with transcriptome sequencing and GO enrichment analysis, five genes related to IAA metabolism were identified to be differentially expressed between G1+TYC and TYC during 30-60 DAF, which might serve as key candidate genes determining the fruit enlargement of G1+TYC.【Conclusion】This study revealed that enlargement of juice sac volume and elevated IAA content in pulp at early stage of fruit development were closely associated with fruit enlargement of G1+TYC. Moreover, candidate genes related to fruit enlargement in G1+TYC were identified based on transcriptome analysis, providing valuable genetic resources for citrus fruit size regulating mechanism research and further genetic improvement.

Key words: Citrus, cybrid, fruit size, endogenous hormones, transcriptome

Table 1

Mass spectrometry parameters of different plant endogenous hormones"

激素种类
Hormones type		 保留时间
Retention time (min)		 母离子
Parent ions (m·z-1)		 子离子
Sub ion(m·z-1)		 离子模式
Ionic mode
吲哚-3-乙酸 IAA 8.6 174 128 负离子Negative
赤霉素 GA3 7.7 345 239 负离子Negative
玉米素 Zeatin 4.8 220 136 正离子Positive
脱落酸 ABA 8.9 263 153 负离子Negative

Table 2

Sequence of primers"

引物名称 Primer name 正向引物 Forward primer (5′-3′) 反向引物 Reverse primer (5′-3′)
核基因组引物
Nuclear genome primer		 TAA15 GAAAGGGTTACTTGACCAGGC CTTCCCAGCTGCACAAGC
线粒体基因组引物
Mitochondrial genome primer		 G1 ACGCTTTGGTTAGGCTTGG GGCTCGAATGCCTTTACG
HB CCCGCCCTTAGGAGATTG TCCCTCGGACTCGGAAAG
叶绿体基因组引物
Chloroplast genome primer		 M ACTCGATATTTCAACCCATCC CACAATTCCGATCCAGAGC
NTCP9 CTTCCAAGCTAACGATGC CTGTCCTATCCATTAGACAATG

Fig. 1

Identification of genetic origin of nuclear and cytoplasmic genomes of the G1+TYC A: Nuclear genome (TAA15); B: Mitochondrial genome (left: G1, right: HB); C: Chloroplast genome (left: M, right: NTCP9). M: DNA marker; 1: Taoye sweet orange; 2: Guoqing No.1 Satsuma mandarin; 3-8: Satsuma mandarin+Taoye sweet orange somatic cybrid"

Fig. 2

Fruits of G1+TYC and TYC at different developmental stages and their fruit growth curves A: Fruits of G1+TYC and TYC collected at 15-240 days after flower; B: Transverse diameter growth curve of G1+TYC and TYC; C: Longitudinal diameter growth curve of G1+TYC and TYC. DAF: Days after flower. *, ** indicate significant differences at P<0.05 and P<0.01, respectively. The same as below"

Fig.3

Phenotypic comparison of fruit size, weight and other morphological traits between G1+TYC and TYC A: Mature fruits of G1+TYC and TYC; B: Mature juice sacs of G1+TYC and TYC; C: Phenotypic comparison of other fruit morphological traits between G1+TYC and TYC"

Fig. 4

Statistics of soluble sugar and organic acid contents in the pulp of G1+TYC and its mesophyll parent TYC at different developmental stages in 2023"

Fig. 5

Determination of plant hormone content in the pulp of G1+TYC and TYC during 2022 and 2023"

Fig. 6

Pathway enrichment of differentially expressed genes between G1+TYC and TYC and expression of candidate genes A: Enrichment of differentially expressed gene (DEGs) pathways between G1+TYC and TYC at 60 days after flower; B: Clustering heatmap of TPM values for five DEGs in the IAA pathway in the pulp of G1+TYC and TYC at 30 days and 60 days after flower"

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