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作物学报 ›› 2026, Vol. 52 ›› Issue (6): 1658-1668.doi: 10.3724/SP.J.1006.2026.55069

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

夜间复合体关键基因AhLUX1正向调控花生结瘤的功能研究

陆一楚**(), 李振影**(), 买春海, 赵晓蕊, 王利祥*()   

  1. 山西农业大学农学院 / 山西省后稷实验室, 山西太谷030801
  • 收稿日期:2025-11-18 接受日期:2026-03-16 出版日期:2026-06-12 网络出版日期:2026-03-31
  • 通讯作者: * 王利祥, E-mail: lxwang@sxau.edu.cn
  • 作者简介:陆一楚, E-mail: luyichu1@sxau.edu.cn;李振影, E-mail: lizhenying@sxau.edu.cn

    **同等贡献

  • 基金资助:
    山西省科技重大专项揭榜挂帅项目(202101140601027);山西农业大学高层次人才启动经费(2021xG003);山西农业大学高层次人才启动经费(2022xG0014)

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 Published:2026-06-12 Published online: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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摘要：

豆科植物结瘤固氮是一个高度耗能的生物学过程。对豆科植物而言, 光信号不仅调控光合作用、形态建成和生长发育等过程, 还对其与根瘤菌互作中的结瘤和固氮过程起关键调控作用。LUX是夜间复合体的核心组分, 该蛋白能够在恒定光照和黑暗条件下保障多种信号的有序输出。与大豆等豆科植物不同, 花生主要通过裂缝入侵完成侵染, 关于AhLUX1在花生结瘤中的调控作用目前尚不清楚。本研究表明, AhLUX1能显著响应根瘤菌的侵染, 并主要在根尖、侧根原基、发育中的根瘤和成熟根瘤中表达。亚细胞定位结果表明, AhLUX1定位于细胞核中。功能研究表明, AhLUX1过表达(OE-AhLUX1)显著增加根瘤数, 沉默(RNAi-AhLUX1)或敲除(CR-AhLUX1)则显著抑制花生的根瘤数, 证明该基因是花生结瘤的正调控因子。为了初步解析AhLUX1调控花生结瘤的机理, 本研究进一步对AhLUX1过表达和沉默株系根中的结瘤标记基因进行检测。结果表明, AhLUX1可能通过调控结瘤关键基因的表达影响花生结瘤。本研究通过解析AhLUX1对花生结瘤的影响, 将光信号调控因子与地下结瘤固氮这2个生物学过程联系起来, 为通过调控光信号通路培育高效结瘤固氮的花生新品种提供了关键的基因资源。

关键词: 花生, AhLUX1基因, 夜间复合体, 结瘤, 共生固氮

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

附表1

本研究所用引物列表"

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

图1

LUXs系统发育树与序列比对 图A为LUXs系统发育树。各分支节点上的数字代表自展值。图B为不同物种LUX蛋白的序列比对。序列上方的数字表示氨基酸残基的位置。下方的黑色横线标记了花生与其他物种间存在差异的重要残基, 下方的蓝色横线标记了LUXs的MYB保守结构域。Ps: 豌豆; Vf: 蚕豆; Ah: 花生; Gm: 大豆; Pv: 菜豆; Vu: 豇豆。"

图2

AhLUX1基因的表达模式分析 图A为AhLUX1在根瘤中的相对表达量。图B为接菌后0-24 h AhLUX1的表达量(以接种后0 h的表达水平为对照)。图C为接菌后1-28 d AhLUX1的表达量(以接种后1 d的表达水平为对照)。图D为AhLUX1在花生不同组织中的表达量。所示数据为平均值±标准误。显著性使用t检验分析。***表示P < 0.001; **表示P < 0.01; *表示P < 0.05。"

图3

pAhLUX1:GUS组织化学染色分析 图A为接种根瘤菌1 d后的根系。图B-D为接种根瘤菌3、6和9 d后的根系。图E为接种根瘤菌14 d后的根系。图F为接种根瘤菌28 d后的根系。比例尺= 1 mm。"

附图1

AhLUX1的亚细胞定位预测"

图4

AhLUX1的亚细胞定位"

图5

AhLUX1调控花生结瘤表型分析 图A为过表达空载体对照(EV1)和OE-AhLUX1的阳性根中AhLUX1的RT-qPCR分析。图B为EV1和OE-AhLUX1株系的毛状根对比图。图C为EV1和OE-AhLUX1株系的毛状根根瘤数目比较。图D为基因沉默空载体对照(EV2)和RNAi-AhLUX1的阳性根中AhLUX1的RT-qPCR分析。图E为EV2和RNAi-AhLUX1株系的毛状根对比图。图F为EV2和RNAi-AhLUX1株系的毛状根根瘤数目比较。图G为基因编辑空载体对照(EV3)和CR-AhLUX1株系的毛状根对比图。图H为EV3和CR-AhLUX1株系的毛状根根瘤数目比较。比例尺= 5 mm。所示数据为平均值±标准误。显著性使用t检验分析。***表示P < 0.001。"

图6

AhLUX1敲除类型分析 红色字体: 突变碱基; 黑色虚线: 省略部分碱基。"

附表2

结瘤标记基因登录号"

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

图7

OE-AhLUX1和RNAi-AhLUX1根系中结瘤标记基因相对表达量分析 图A为OE-AhLUX1根系中结瘤标记基因的相对表达量。图B为RNAi-AhLUX1根系中结瘤标记基因的相对表达量。所示数据为平均值±标准误。显著性使用t检验分析。***表示P < 0.001; **表示P < 0.01; *表示P < 0.05; ns表示差异不显著。缩写同图5。"

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