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Acta Agronomica Sinica ›› 2026, Vol. 52 ›› Issue (6): 1658-1668.doi: 10.3724/SP.J.1006.2026.55069

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

Positively regulating role of the key evening complex gene AhLUX1 in peanut nodulation

Lu Yi-Chu**(), Li Zhen-Ying**(), Mai Chun-Hai, Zhao Xiao-Rui, Wang Li-Xiang*()   

  1. College of Agriculture, Shanxi Agricultural University / Shanxi Houji Laboratory, Taigu 030801, Shanxi, China
  • Received:2025-11-18 Accepted:2026-03-16 Online:2026-06-12 Published:2026-03-31
  • Contact: * Wang Li-Xiang, E-mail: lxwang@sxau.edu.cn
  • About author:

    **Contributed equally to this work

  • Supported by:
    Major Special Science and Technology Projects in Shanxi Province(202101140601027);Startup Funds for High-Level Talents of Shanxi Agricultural University(2021xG003);Startup Funds for High-Level Talents of Shanxi Agricultural University(2022xG0014)

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Abstract:

Biological nitrogen fixation via nodulation in legumes is highly energy intensive. In legumes, light signals not only regulate photosynthesis, morphogenesis, and growth, but also play key roles in controlling nodulation and nitrogen fixation during interactions with rhizobia. LUX (lux arrhythmo) is a core component of the evening complex (EC), which ensures accurate temporal output of multiple signals under light-dark cycles. Unlike soybean and many other legumes, peanut primarily uses a crack-entry mode of rhizobial invasion, and the role of AhLUX1 in peanut nodulation has remained unclear. Here, this study shows that AhLUX1 is strongly induced by rhizobial infection and is predominantly expressed in root tips, lateral root primordia, developing nodules, and mature nodules. Subcellular localization analysis indicates that AhLUX1 is localized in the nucleus. Functional analyses show that overexpression of AhLUX1 (OE-AhLUX1) significantly increases nodule number, whereas silencing (RNAi-AhLUX1) or knockout (CR-AhLUX1) markedly reduces nodulation, demonstrating that AhLUX1 is a positive regulator of nodulation in peanut. To clarify the underlying mechanism, the expression of nodulation marker genes in AhLUX1-overexpressing and AhLUX1-silenced roots. The results suggest that AhLUX1 modulates peanut nodulation by regulating the expression of key nodulation genes. Together, these findings link a light-signaling component to belowground nodulation and nitrogen fixation and provide a genetic resource for breeding high-efficiency peanut varieties through manipulation of the light-signaling pathway.

Key words: peanut, AhLUX1, evening complex, nodulation, symbiotic nitrogen fixation

Table S1

List of primers used in this study"

引物名称
Primer name		 引物序列
Primer sequence (5'-3')
AhLUX1-F ATGGGTGAGGAAGTTAGGATG
AhLUX1-R AGTTTCCTTTGATTGTCTTCCAAC
OE-AhLUX1-F ttgatgtgattacagtctagaATGGGTGAGGAAGTTAGGATGACC
OE-AhLUX1-R gttaattaacccgctggtaccAGTTTCCTTTGATTGTCTTCCAACC
AhLUX1-207-F GGGGACAAGTTTGTACAAAAAAGCAGGCTTCATGGGTGAGGAAGTTAGGATG
AhLUX1-207-R GGGGACCACTTTGTACAAGAAAGCTGGGTCAGTTTCCTTTGATTGTCTTCCAAC
AhLUX1-1300-F CCAAATCGACTCTAGAAAGCTTATGGGTGAGGAAGTTAGGATG
AhLUX1-1300-R CCACTAGTATTTAAATGTCGACGAGTTTCCTTTGATTGTCTTCCAAC
AhLUX1-BSF0 ATATATGGTCTCGATTGGAGTCGCCGACTGGGAATTGTT
AhLUX1-F0 TGGAGTCGCCGACTGGGAATTGTTTTAGAGCTAGAAATAGC
AhLUX1-R0 AACCGTCTTGGAAGGGGTTGAGCAATCTCTTAGTCGACTCTAC
AhLUX1-BSR0 ATTATTGGTCTCGAAACCGTCTTGGAAGGGGTTGAGCAA
AhLUX1-Test-F ATGGGTGAGGAAGTTAGGATGAC
AhLUX1-Test-R AATCCGCTTCCTCCGCACAATC
pAhLUX1-F GGCTCTACAGAGGTGCAAGG
pAhLUX1-R CCGACTACCAAGACGACGAC
pAhLUX1-1391-F TGGGCCCGGCGCGCCAAGCTTGGCTCTACAGAGGTGCAAGGAG
pAhLUX1-1391-R TCTTAGAATTCCCGGGGATCCGTCGTCGTCTTGGTAGTCGGTC
AhActin-F GAAGTTCTCTTCCAACCATCC
AhActin-R TTTCCTTGCTCATTCTGTCCG
qRT-AhLUX1-F CCACATATTGGTGTGCCTGC
qRT-AhLUX1-R TCCAACCAACTTTACCCTTTCT
qRT-AhLUX2-F CCACCACATATTGGAGTGCC
qRT-AhLUX2-R CCAACCAACTTTACCCTTTCTTTGA
qRT-AhLUX1-K7-F TTGGTGGTCATGGACTCATGG
qRT-AhLUX1-K7-R TGGCATTAGGTGCTCCTCTTT
qRT-AhNIN-F GGTACCTATGAGGAGGAGATCAGCC
qRT-AhNIN-R AGCATTAGAGAGATGATCAAGTTGTGG
qRT-AhENOD40-F GGCAAAAATCCATCCATGGTTC
qRT-AhENOD40-R GCCTTTTTGTGACTTGCCGG
qRT-AhEFD-F CAATTCCCAATTGGTGCTGGAACAATTGC
qRT-AhEFD-R GACCCAATTGAGCAAAGTTCAATTGATCCAC
qRT-AhCCamK-F GCCAAAGATGTTCAAATGGACCCT
qRT-AhCCamK-R CCTGAGACTTTCAATTTCCTCTTC
qRT-AhHK-F GAGGTCTTAAAGCCATTGTGGTTGATGG
qRT-AhHK-R CAACTGCTTGATAGAGATTCTCTTC
qRT-AhSYMRK-F CAGATCAGCTTCTGCAAGTGAAGG
qRT-AhSYMRK-R GAAATGTGCCCCTTATCAGATACACTTTG
qRT-AhREM-F GACTAAAATAGACAACAGGGC
qRT-AhREM-R GCAGTAAAGCATGAAAGAATTTTC
qRT-AhELF-F AAGCTTCCCTGGCAAAGCTCAA
qRT-AhELF-R TTCCTCAGCTGCCTTCTTATCC

Fig. 1

Phylogenetic analysis and sequence alignment of LUXs A: phylogenetic tree of LUXs. Numbers at internal nodes indicate bootstrap support values. B: amino acid sequence alignment of LUXs from different species. Numbers above the sequences indicate amino acid positions. Black bars indicate key residues that differ between peanut and other species, and the blue bar indicates the conserved MYB domain of LUXs. Ps: Pisum sativum; Vf: Vicia faba; Ah: Arachis hypogaea; Gm: Glycine max; Pv: Phaseolus vulgaris; Vu: Vigna unguiculata."

Fig. 2

Expression pattern analysis of AhLUX1 A: relative expression level of AhLUX1 in nodules. B: relative expression level of AhLUX1 from 0 to 24 h after rhizobial inoculation (normalized to 0 h). C: relative expression level of AhLUX1 from 1 to 28 d after rhizobial inoculation (normalized to 1 d). D: AhLUX1 expression in different peanut tissues. Data are means ± SE. Significance is analyzed by t-tests. ***, P < 0.001; **, P < 0.01; *, P < 0.05."

Fig. 3

pAhLUX1:GUS histochemical staining analysis A: roots of 1 d after inoculation with rhizobacteria. B-D: roots of 3, 6, and 9 d after inoculation. E: roots of 14 d after inoculation. F: roots of 28 d after inoculation. Scale bars = 1 mm."

Fig. S1

Prediction of subcellular localization of AhLUX1"

Fig. 4

Subcellular localization of AhLUX1"

Fig. 5

Phenotypic analysis of AhLUX1 function in peanut nodulation A: RT-qPCR analysis of AhLUX1 expression in empty vector control for overexpression (EV1) and OE-AhLUX1. B: representative images of hairy roots from EV1 and OE-AhLUX1. C: nodule numbers in EV1 and OE-AhLUX1 hairy roots. D: RT-qPCR analysis of AhLUX1 expression in empty vector control for gene silencing (EV2) and RNAi-AhLUX1. E: representative images of hairy roots from EV2 and RNAi-AhLUX1. F: nodule numbers in EV2 and RNAi-AhLUX1 hairy roots. G: representative images of hairy roots from empty vector control for gene editing (EV3) and CR-AhLUX1. H: nodule numbers in EV3 and CR-AhLUX1 hairy roots. Scale bar = 5 mm. Data are means ± SE. Significance is analyzed by t-tests. ***, P < 0.001."

Fig. 6

Analysis of AhLUX1 knockout types Red font: mutated bases; black dashed lines: omitted bases."

Table S2

Accession numbers of nodulation marker genes"

基因名称
Gene name		 登录号
Accession number		 参考来源
Reference source
AhNIN KT820787 Das, et al. [25]
AhENOD40 KY401676
AhSYMREM KT988038
AhEFD KT962993
AhCCamK EU395429
AhHK KT153625
AhSYMRK FJ969396.2

Fig. 7

Expression analysis of nodulation marker genes in OE-AhLUX1 and RNAi-AhLUX1 roots A: relative expression of nodulation marker genes in roots of OE-AhLUX1. B: relative expression of nodulation marker genes in roots of RNAi-AhLUX1. Data are means ± SE. Significance is analyzed by t-tests. ***, P < 0.001; **, P < 0.01; *, P < 0.05; ns, no significant difference. Abbreviations are the same as those given in Fig. 5."

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