Scientia Agricultura Sinica ›› 2026, Vol. 59 ›› Issue (1): 1-16.doi: 10.3864/j.issn.0578-1752.2026.01.001

• CROP GENETICS & BREEDING·GERMPLASM RESOURCES·MOLECULAR GENETICS • Previous Articles     Next Articles

Functions of ABC Transporter OsARG1 in Rice Heading Date Regulation

WANG ZhongNi(), LEI Yue, LI JiaLi, GONG YanLong, ZHU SuSong*()   

  1. Rice Research Institute, Guizhou Academy of Agricultural Sciences, Guiyang 550006
  • Received:2025-06-18 Accepted:2025-08-08 Online:2026-01-01 Published:2026-01-07
  • Contact: ZHU SuSong

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Abstract:

【Objective】Heading date is a critical agronomic trait influencing rice yield and quality, regulated by complex networks involving histone-modifying enzymes, transcription factors, protein kinases, florigens, and phytochromes. While ATP-binding cassette (ABC) transporters are known for their roles in substrate transport, their functions in heading date regulation remain unclear. This study investigates the role of the ABC transporter gene OsARG1 in the regulation of rice heading date, which will provide evidence for enriching the heading date regulation network.【Method】A comparative analysis was conducted between wild-type Nipponbare and the osarg1 mutant. Key agronomic traits, including heading date, plant height, tiller number, and panicle length, were assessed. Chlorophyll contents in leaf of Nipponbare (WT), albino leaf (WL), yellow-green leaf (YL) and green leaf (GL) of osarg1 were measured. Metal element contents such as cobalt, nickel, calcium, magnesium and iron in WT, WL, YL and GL were determined by ICP-MS. Hormone profiling, transcriptome sequencing, and integrative analysis were performed on WT, WL and GL to explore the regulatory function of OsARG1.【Result】The osarg1 mutant exhibited an earlier heading date than wild-type in both Guiyang and Changchun. It also showed reduced plant height, tiller number, and panicle length. Chlorophyll levels in WL and YL were significantly lower, accompanied by disrupted metal element homeostasis. Hormonal analysis revealed elevated levels of gibberellins, auxin-related, and cytokinin-related hormones in GL and WL, particularly in WL. Transcriptome analysis identified 2 001 and 6 555 differentially expressed genes (DEGs) in GL_vs_WT and WL_vs_WT comparisons, respectively. Over 20 heading date-associated genes, including Hd3a, OsMADS14, and chromatin methyltransferase genes, were differentially expressed. GO and KEGG analyses of DEGs from GL vs WT comparison highlighted enrichment in pathways related to metabolism, development, and environmental responses. Integrated transcriptomic and hormonal analysis suggested that OsARG1 may influence gibberellin and cytokinin levels by modulating diterpene and zeatin metabolism and hormone signaling pathways. Expression levels of the selected genes by qRT-PCR were consistent with the transcriptome data, validating the transcriptomic findings. 【Conclusion】The osarg1 mutant heads earlier than the wild-type, with OsARG1 likely regulating heading date through modulating the expression of heading date related genes. Additionally, OsARG1 plays roles in maintaining chlorophyll content and metal element (such as nickel, iron, and magnesium) balance in rice leaves.

Key words: rice, heading date, OsARG1, metal element content, phytohormone, transcriptome sequencing

Table 1

Primers for qRT-PCR"

类别
Classification		 基因
Gene		 基因ID
Gene ID		 正向引物
Forward primer (5′-3′)		 反向引物
Reverse primer (5′-3′)
抽穗期相关基因
Heading date related genes		 Hd3a LOC4340185 AGCCCAAGTGACCCTAACCT GTTGTAGAGCTCGGCGAAGT
OsMADS14 LOC4334140 TGAAGCGGATCGAGAACA CATGAGTCGGTGGCGTAC
激素信号相关基因
Hormone signaling related genes		 OsEBF2 LOC4328647 TGGCATCAGCAAAGCACC ATGTCGCACCACCAGAGC
GID1 LOC4338764 GCCCATCCTTGAGTTCCTGAC ACACCACGACGCCCTTGCTC
激素代谢相关基因
Hormone metabolism related genes		 OsIPT7 LOC4339535 GCGAGGATACGAGGATGGTGG CAGCGTCCACCATGTCGTCC
ZEATIN LOC4336630 CGCACCCGATGTTCAAAGA GCCGAATGACACGTAGAGGA
CPS4 LOC4335090 TCCAACCCATCCTCTGTCA GGCCCACTTGTCCTTCATT
GA20ox LOC4336431 AGCAGGCAAGGCTGTTCCG GTGAGGTCTCCAAAGTCGCAAT
内参基因
Internal reference gene		 ACTIN1 LOC4333919 TGCTATGTACGTCGCCATCCAG AATGAGTAACCACGCTCCGTCA

Fig. 1

Heading dates and agronomic traits of WT and osarg1 mutant A: Heading dates of WT and osarg1 mutant in Changchun, Jilin province; B: Heading dates of WT and osarg1 mutant in Guiyang, Guizhou province; C: Statistics of heading date of WT and osarg1 in Changchun (CC) and Guiyang (GY); D-F: Panicle length, grain length and grain width of WT and osarg1; G: osarg1 mutant produced higher tillers. The arrow shows the higher tiller in osarg1 after heading; H: Leaf color. It shows the leaf of WT, yellow-green leaf (YL), albino leaf (WL) and green leaf (GL) of osarg1 from left to right. bar=1 cm"

Table 2

Agronomic traits of wild type and osarg1 in Guiyang"

名称
Name		 株高
PH (cm)		 分蘖数
TN		 穗长
PL (cm)		 穗粒数
SPP		 结实率
SR (%)		 千粒重
TGW (g)		 粒长
SL (mm)		 粒宽
SW (mm)
WT 73.8±3.2 11.7±2.4 17.8±1.2 85.7±17.2 94.3±4.3 25.5±0.6 7.6±0.2 3.7±0.1
osarg1 40.0±4.0 5.4±1.7 11.3±1.6 27.3±5.5 70.9±1.7 20.6±0.1 7.2±0.1 3.2±0.1

Fig. 2

Chlorophyll content in different color leaves of rice osarg1 mutant A: Chlorophyll a contents; B: Chlorophyll b contents; C: Total chlorophyll contents. WT: Leaves of wildtype; WL: Albino leaf; YL: Yellow-green leaf; GL: Green leaf. *: Significant difference at P<0.05 level; **: Significant difference at P<0.01 level. The same as below"

Table 3

Metal element contents in leaves of different colors (μg·g-1)"

名称Name Na Mg K Ca Mn Fe Co Ni Cu Zn As Se Rb Sr Pb
野生型WT 68.70 266.41 4020.38 3.75 166.21 227.17 0.06 0.11 3.75 11.58 0.14 0.24 0.73 6.84 0.17
白化叶WL 86.08 499.72 8058.26 3.16 162.70 73.35 0.07 - 3.16 12.81 6.44 0.18 3.13 3.97 0.31
黄绿叶YL 40.76 305.40 4155.11 1.74 231.35 129.06 0.04 0.03 1.74 7.33 0.11 0.14 1.34 6.91 0.17
绿色叶GL 165.38 769.00 3892.28 5.13 711.47 219.34 0.08 0.49 5.13 10.28 0.61 0.23 0.94 27.47 0.62

Table 4

Hormone contents in leaves of different colors (ng·g-1)"

激素
Hormone		 名称
Name		 激素含量 Hormone contents
野生型
WT1		 野生型
WT2		 野生型
WT3		 白化叶
WL1		 白化叶
WL2		 白化叶
WL3		 绿色叶
GL1		 绿色叶
GL2		 绿色叶
GL3
赤霉素
Gibberellin		 GA3 0.00 0.00 0.00 20.53 20.82 19.57 10.48 10.40 9.82
GA1 14.53 21.81 19.69 112.36 112.36 98.82 26.55 29.56 30.92
生长素
Auxin		 ICA 18.41 22.11 26.29 138.08 106.76 147.49 57.25 37.83 40.56
MEIAA 0.00 0.00 0.00 31.02 33.18 33.39 5.15 4.16 7.01
3-IPA 0.00 0.00 0.00 142.97 0.00 0.00 0.00 0.00 0.00
tIAA 5.43 5.92 4.34 16.12 17.40 17.68 4.92 5.08 6.55
IAA 43.50 50.14 52.72 437.59 399.94 434.50 60.11 53.33 58.79
细胞分裂素
Cytokinin		 IP 0.00 0.00 0.00 4.41 0.00 0.00 0.00 0.00 0.00
cZ 109.15 92.10 86.79 334.13 288.51 251.73 9.88 9.87 11.77
Dx 120.69 124.34 118.61 112.59 97.99 103.45 212.56 183.52 190.03
Dh-Z 104.88 87.15 103.32 17.77 18.11 16.71 31.41 33.13 37.42
IPA 203.75 202.39 194.53 89.58 107.28 119.08 106.11 122.79 122.02
水杨酸
Salicylic acid		 MESA 0.00 0.00 0.00 2043.71 1945.56 1960.40 0.00 0.00 0.00
SA 262841.1 208707.3 226887.3 148707.0 143286.1 191767.7 140563.7 151651.5 170172.6
茉莉酸
Jasmonic acid		 H2JA 2.36 1.83 2.76 3.06 2.60 3.11 2.75 2.88 3.03
JA 93.82 83.59 76.27 63.44 50.83 59.65 377.76 393.99 492.27
脱落酸
Abscisic acid		 ABA 57.47 47.28 45.39 150.24 120.25 97.19 55.13 49.21 36.48
乙烯
Ethylene		 ACC 838.67 778.14 822.08 5323.73 5453.37 5235.46 3885.59 3295.39 3467.53

Fig. 3

Contents of common differential hormone in WL and GL of osarg1 mutant A: Gibberellin; B: Auxin; C: Cytokinin; D: ACC. Different lowercase letters represent significant difference"

Table 5

Quality and genome comparison of transcriptome sequencing data"

样品名称Samples 过滤后总数据量
Clean bases		 GC含量
GC content (%)		 ≥Q30
(%)		 序列总数
Total clean reads		 总比对（比对率）
Total mapped (Mapping ratio, %)		 唯一比对（比对率）
Uniquely mapped (Mapping ratio, %)		 多方比对（比对率）
Multiple mapped (Mapping ratio, %)
绿色叶GL1 7302373276 50.47 91.07 48819940 46388953 (95.02) 44247888 (90.63) 2141065 (4.39)
绿色叶GL2 7434294208 50.05 91.25 49705204 47042791 (94.64) 43805600 (88.13) 3237191 (6.51)
绿色叶GL3 7271079057 49.14 91.54 48606466 45884410 (94.40) 43524507 (89.54) 2359903 (4.86)
白化叶WL1 6830948082 50.15 92.11 45672892 43865510 (96.04) 42366504 (92.76) 1499006 (3.28)
白化叶WL2 7148140757 49.58 91.73 47793474 45823105 (95.88) 43672117 (91.38) 2150988 (4.50)
白化叶WL3 7944166308 49.72 97.67 53103966 52271280 (98.43) 49985049 (94.13) 2286231 (4.31)
野生型WT1 6591969488 53.31 93.37 44115034 41355884 (93.75) 37259893 (84.46) 4095991 (9.28)
野生型WT2 6763621452 50.12 92.11 45189738 43024853 (95.21) 41764281 (92.42) 1260572 (2.79)
野生型WT3 8854258271 53.17 95.02 59169498 56245607 (95.06) 50550176 (85.43) 5695431 (9.63)

Fig. 4

Transcriptome analysis of heading date related genes in osarg1 A: Up-regulated and down-regulated DEGs in WL and GL; B: Heat map of expression levels of heading date related genes in WT and GL"

Fig. 5

Enrichment analysis of differential expressed genes in GL_vs_WT A: GO enrichment analysis of differential expressed genes; B: Enriched chord map of top 10 GO items in biological process. Lines between genes and GO entries represent enrichment relationship between them; C: KEGG enrichment analysis of differential expressed genes in biological process"

Fig. 6

KEGG enrichment analysis of hormone content changes in green leaf of osarg1 A: KEGG enrichment bubble plot of differential genes/metabolites. Bubble shapes represent different omics, circles represent transcriptomes, and triangles represent metabolomes; B-C: KEGG pathway diagrams of gibberellin and cytokinin, respectively. Boxes indicate genes, and circles indicate metabolites. Red indicates upregulated genes/metabolites, green indicates downregulated genes/metabolites, blue indicates upregulated and downregulated genes/metabolites. Gene expression levels are represented by heat map. The left and right three squares in heat map are genes expression levels in WT and GL, respectively. The solid line represents specific reaction, while the dashed line represents omitted multiple reactions"

Fig. 7

Enrichment analysis of differential expressed genes in WL and GL of osarg1 A:Venn diagrams of differential expressed genes in WT, GL and WL; B: KEGG analysis of 3 497 differential expressed genes"

Fig. 8

Relative expression levels of differential expressed genes in osarg1 A: FPKM values of DEGs from RNA-seq; B: Relative expression levels of DEGs by qRT-PCR"

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