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作物学报

作物学报 ›› 2026, Vol. 52 ›› Issue (1): 28-43.doi: 10.3724/SP.J.1006.2026.51069

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

藜麦RopGEF家族基因的鉴定及表达模式分析

景秀清1,2,3,*(), 蔡永朵1, 邓宁1, 赵晓东1,2, 翟飞红1,2, 曾群4   

  1. 1太原师范学院生物科学与技术学院, 山西晋中 030619
    2太原师范学院汾河流域地表过程与资源生态安全山西省重点实验室, 山西晋中 030619
    3山西大学生命科学学院, 山西太原 030006
    4山西中医药大学基础医学院, 山西晋中 030619
  • 收稿日期:2025-07-22 接受日期:2025-10-30 出版日期:2026-01-12 网络出版日期:2025-11-10
  • 通讯作者: *景秀清, E-mail: xiuqingjing@tynu.edu.cn
  • 基金资助:
    山西省基础研究计划(自由探索类)项目(202203021212186);太原师范学院2024年度研究生教育创新项目(SYYJSYC-2431);太原师范学院大学生创新创业训练项目(CXCY25104)

Identification and expression pattern analysis of RopGEF family genes in Chenopodium quinoa

Jing Xiu-Qing1,2,3,*(), Cai Yong-Duo1, Deng Ning1, Zhao Xiao-Dong1,2, Zhai Fei-Hong1,2, Zeng Qun4   

  1. 1College of Biological Sciences and Technology, Taiyuan Normal University, Jinzhong 030619, Shanxi, China
    2Shanxi Key Laboratory of Earth Surface Processes and Resource Ecology Security in Fenhe River Basin, Taiyuan Normal University, Jinzhong 030619, Shanxi, China
    3College of Life Science, Shanxi University, Taiyuan 030006, Shanxi, China
    4School of Basic Medical, Shanxi University of Chinese Medicine, Jinzhong 030619, Shanxi, China
  • Received:2025-07-22 Accepted:2025-10-30 Published:2026-01-12 Published online:2025-11-10
  • Contact: *E-mail: xiuqingjing@tynu.edu.cn
  • Supported by:
    Fundamental Research Programs of Shanxi Province(202203021212186);Graduate Education and Innovation Projects of Taiyuan Normal University(SYYJSYC-2431);College Student Innovation and Entrepreneurship Training Program of Taiyuan Normal University(CXCY25104)

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摘要：

ROP鸟嘌呤核苷酸交换因子(Rho-related GTPase of plants guanine nucleotide-exchange factors, RopGEFs)介导的ROP信号转导在植物细胞信号通路中起着关键作用。本研究利用生物信息学分析方法, 从藜麦基因组中共鉴定到7个RopGEF家族基因(分布于6条染色体上)。基于系统进化关系和结构特征, 将其与拟南芥、水稻等6种作物的共90个RopGEFs分为4个亚家族, 其中CqRopGEF5与AtRopGEF1、OsRopGEF1亲缘关系较近。结构分析显示, 同亚家族CqRopGEFs的外显子-内含子分布、蛋白Motifs组成及二级/3D结构均具保守性。实时荧光定量聚合酶链式反应(quantitative real-time polymerase chain reaction, qRT-PCR)结果表明, 多数CqRopGEFs在种子萌发期高表达, 且幼苗根中表达量高于茎、叶; 其表达受外源ABA和非生物胁迫显著诱导, 如ABA处理下CqRopGEF2/3/4/7表达先升后降, CqRopGEF7在冷、热胁迫后急剧下调。综上, CqRopGEFs家族结构进化保守, 可能参与藜麦的生长发育、ABA信号通路及非生物胁迫响应。

关键词: 藜麦, RopGEFs, 系统进化分析, 基因表达, 非生物胁迫

Abstract:

In plants, RopGEF-mediated ROP signaling plays a crucial role in cellular signaling pathways. In this study, seven RopGEF family members were identified in Chenopodium quinoa through bioinformatics analysis and were found to be distributed across six chromosomes. Based on phylogenetic relationships and structural characteristics, 90 RopGEFs from six crop species, including Arabidopsis thaliana and rice, were classified into four subfamilies. Evolutionary analysis revealed that CqRopGEF5 is closely related to AtRopGEF1 and OsRopGEF1. Structural analysis indicated that the exon-intron organization, protein motif composition, and secondary and three-dimensional structures of CqRopGEFs are highly conserved. qRT-PCR analysis showed that most CqRopGEFs were highly expressed during seed germination, with expression levels in seedling roots being higher than in stems and leaves. Their expression was significantly induced by exogenous abscisic acid (ABA) and abiotic stresses. For instance, the expression of CqRopGEF2, CqRopGEF3, CqRopGEF4, and CqRopGEF7 initially increased and then decreased under ABA treatment, while CqRopGEF7 was strongly downregulated under both cold and heat stress. In conclusion, the CqRopGEF gene family exhibits conserved evolutionary structure and may play important roles in quinoa growth and development, ABA signaling, and responses to abiotic stress.

Key words: Chenopodium quinoa, RopGEFs, phylogenetic analysis, gene expression, abiotic stresses

表1

qRT-PCR引物序列"

引物名称
Primer name		 引物序列
Primer sequence (5'-3')
CqActin-F CCCCTGCTATGTATGTTGCAATTC
CqActin-R AGTGGTCTGTTAGGTCACGACCAG
CqRopGEF1-F GTCAGAACCGGATAGGAATGACA
CqRopGEF1-R TTCTAAGATTGCTTGGCCGACG
CqRopGEF2-F GTCAAGTGCTGAGCGTAGAA
CqRopGEF2-R AGAGTAGGCGAAACAGAGGA
CqRopGEF3-F TGATTCAGGGCTTGATGAATTAG
CqRopGEF3-R CTGCAGTACATACACCTTTCCC
CqRopGEF4-F AAGATCACTCGAGCAATCCCAA
CqRopGEF4-R CTTATATGTGGGTTGCCTATGG
CqRopGEF5-F TCTCCCGATCCAAAAACGCA
CqRopGEF5-R ATCTAGGGCTTCCTCCAGCA
CqRopGEF6-F CTGCTCGACATTGACAAGAGAC
CqRopGEF6-R AACAAGTGCTTTCTCTCTCGC
CqRopGEF7-F GAGAAGTCCAACAGCTCGACA
CqRopGEF7-R AGGGAAGTTAAAACTGTTGAGGA

图1

藜麦RopGEF家族基因的染色体定位"

表2

藜麦RopGEF家族基因的鉴定"

命名
Name		 基因ID
Gene ID		 染色体定位
Genomic location		 方向
Orientation		 DNA
(bp)		 mRNA
(bp)		 CDS
(bp)		 蛋白质
Protein (aa)
CqRopGEF1 AUR62022562 Chromosome 1A: 2964588-2967787 正向Forward 3200 2059 1344 447
CqRopGEF2 AUR62022802 Chromosome 2A: 55459754-55456379 反向Reverse 3377 2493 1806 601
CqRopGEF3 AUR62004356 Chromosome 2B: 72815241-72818458 正向Forward 3218 2102 1347 448
CqRopGEF4 AUR62017402 Chromosome 5B: 11732273-11725942 反向Reverse 8837 2220 1668 555
CqRopGEF5 AUR62012977 Chromosome 7B: 2293460-2296982 正向Forward 3525 2278 1716 571
CqRopGEF6 AUR62016478 Chromosome 7B: 11948979-11952944 正向Forward 3949 1657 1344 447
CqRopGEF7 AUR62011650 Chromosome 9A: 47835328-47832045 正向Forward 3288 2087 1656 551

表3

藜麦RopGEF家族蛋白质的理化性质分析"

名称
Name		 蛋白质分子量
MV (Da)		 等电点
pI		 电荷
Charge		 不稳定指数
II		 脂肪指数
AI		 总平均亲水性
GRAVY		 主要氨基酸含量
Major amino acids content (%)
CqRopGEF1 50733.64 5.64 -5.0 44.12 69.46 -0.532 Ser (9.8), Leu (9.4), Glu (7.8)
CqRopGEF2 67003.52 8.91 15.5 51.88 82.10 -0.468 Ser (12.0), Leu (10.1), Lys (7.0)
CqRopGEF3 50820.74 5.65 -5.0 47.32 68.86 -0.549 Ser (9.8), Leu (9.4), Glu (8.0)
CqRopGEF4 61739.50 5.42 -8.5 50.76 75.59 -0.447 Ser (13.2), Leu (8.1), Glu (7.6)
CqRopGEF5 63181.72 5.43 -7.0 53.29 79.98 -0.343 Ser (11.0), Leu (10.7), Ala (8.2)
CqRopGEF6 49940.71 5.82 -2.5 44.40 83.18 -0.568 Leu (10.3), Ser (8.7), Lys (8.1)
CqRopGEF7 61880.51 6.72 2.5 46.90 85.35 -0.543 Leu (10.0), Ser (9.1), Lys (8.3)

图2

拟南芥、水稻和藜麦中RopGEF家族成员的系统进化分析"

图3

CqRopGEF家族蛋白质的motif组成"

图4

CqRopGEF家族基因的外显子-内含子组成"

图5

CqRopGEF家族蛋白的二级结构预测"

表4

CqRopGEF家族蛋白质的二级结构比例"

蛋白名称
Protein name		 α-螺旋
Alpha helix		 延伸链
Extended strand		 β-折叠
Beta turn		 无规则卷曲
Random coil
CqRopGEF1 48.55 9.40 6.26 35.79
CqRopGEF2 39.60 11.81 5.16 43.43
CqRopGEF3 49.33 9.38 6.47 34.82
CqRopGEF4 39.46 13.51 5.77 41.26
CqRopGEF5 45.01 11.73 5.78 37.48
CqRopGEF6 50.56 10.29 5.82 33.33
CqRopGEF7 48.46 12.89 6.53 32.12

图6

CqRopGEF家族蛋白质的3D结构模型"

图7

CqRopGEFs的组织特异性表达模式. X轴表示来自藜麦不同组织的不同基因, Y轴显示每个基因的相对表达水平, 以幼苗阶段叶片对照组中对应基因的表达水平作为参考。Seeds at the germination stage: 萌发期的种子; Roots at the seedling stage: 幼苗期的根; Stems at the seedling stage: 幼苗期的茎; Leaves at the seedling stage: 幼苗期的叶。图中误差棒表示3个独立生物重复试验的标准差, 反映了试验数据的变异性。图上小写字母表示组间的统计显著性差异(P < 0.05)。"

附表1

CqRopGEFs的组织特异性相对表达量数据"

藜麦的不同组织
Different tissues of Chenopodium quinoa		 CqRopGEF1 CqRopGEF2 CqRopGEF3 CqRopGEF4 CqRopGEF5 CqRopGEF6 CqRopGEF7
萌发期的种子
Seeds at the
germination stage		 4.878203 2.278607 2.754176 16.890020 2.797100 11.876840 19.897840
3.766052 3.312942 4.805450 15.268320 3.482195 9.355844 16.520080
6.100269 3.838499 6.404784 18.461560 3.013842 5.053957 23.091410
幼苗期的根
Roots at the
seedling stage		 6.807494 1.331894 0.460650 21.580820 3.284094 4.173666 6.313694
3.532024 1.830430 2.551489 20.974230 4.843827 5.355065 6.104382
5.169742 1.199528 0.971060 22.187410 5.968582 6.533347 5.893400
幼苗期的茎
Stems at the
seedling stage		 4.733415 1.770585 1.180153 4.127276 0.713631 0.588344 0.717597
6.430234 1.379614 1.270806 5.117608 2.263214 0.303270 0.691531
1.542182 0.988338 0.902921 4.178092 0.824355 0.451662 0.728712
幼苗期的叶
Leaves at the
seedling stage		 0.518102 1.125639 1.652672 1.051103 1.411523 1.807061 1.370787
0.936619 0.608740 0.535888 0.935297 0.857690 1.133261 0.897113
1.573340 1.642738 1.129119 1.017196 0.826002 0.488311 0.813173

图8

CqRopGEFs响应植物激素ABA的表达模式 X轴代表脱落酸ABA处理组(浓度为100 μmol L-1), Y轴显示各处理组下基因的相对表达量, 以0 h时的基因表达量作为对照进行比较。误差棒代表由3次独立生物学重复计算得出的标准差, 可反映试验数据的变异性。图上小写字母表示各处理组间的显著差异(P < 0.05)。"

附表2

CqRopGEFs响应植物激素ABA的相对表达量数据"

脱落酸诱导时间
ABA induction time (h)		 CqRopGEF1 CqRopGEF2 CqRopGEF3 CqRopGEF4 CqRopGEF5 CqRopGEF6 CqRopGEF7
0 0.383349 0.560141 0.854405 0.606571 0.728405 0.728405 1.062129
1.506241 1.269373 1.345321 1.828694 1.029036 1.029036 0.721514
1.731853 1.406414 0.869982 0.901525 1.334125 1.334125 1.304915
0.5 0.983333 1.102365 1.681996 2.789007 1.298839 9.554574 2.288396
1.524082 1.747551 1.203658 2.703967 1.087367 11.379720 1.333165
2.197235 1.325759 2.439122 1.717055 1.389695 10.233450 0.362952
1.0 3.337438 2.589671 5.702759 3.252600 2.760910 9.061889 4.074006
3.772568 1.520832 4.813831 2.026537 3.317331 10.027570 2.866530
3.308728 1.739145 2.253050 1.514832 2.865699 10.617900 3.495186
3.0 1.137558 3.965328 2.342961 3.991537 0.879867 8.917276 2.692579
1.323412 5.386514 2.337639 5.240955 1.371273 10.102700 4.462155
1.509266 4.675921 2.198124 5.679425 1.445019 9.509880 4.253782
6.0 2.052419 3.843384 2.081516 4.373684 0.209642 18.590750 2.624539
2.699639 3.878513 2.878217 4.111525 0.139735 15.317400 4.317009
4.581864 3.860945 1.946418 3.849441 0.254140 11.211030 1.693145
9.0 2.842707 2.123543 1.361779 2.529633 2.378183 7.451357 1.053495
5.334447 2.349829 2.334819 4.288677 2.990309 9.269848 2.691308
6.244401 1.185006 2.165277 3.323484 2.684246 7.931538 1.479827
12.0 1.089972 1.704662 0.582268 1.517851 0.522919 5.989391 2.404398
1.123806 1.002880 0.458469 1.095538 0.452549 6.101458 0.909693
1.463621 1.428291 0.968643 1.940164 0.657857 5.563212 2.143747

图9

CqRopGEFs响应非生物胁迫的表达模式 X轴代表不同处理组, Y轴显示各处理组下基因的相对表达量, 以0 h时的基因表达量作为对照进行比较。误差线代表由3次独立生物学重复计算得出的标准差, 可反映试验数据的变异性。图上小写字母用于表示各处理组间的显著差异(P < 0.05)。"

附表3

CqRopGEFs响应非生物胁迫的相对表达量数据"

非生物胁迫类型
Abiotic stress types		 诱导时间
Induction time (h)		 CqRopGEF1 CqRopGEF2 CqRopGEF3 CqRopGEF4 CqRopGEF5 CqRopGEF6 CqRopGEF7
盐胁迫
NaCl stress		 0 0.383349 0.560141 0.854405 0.606571 0.728405 0.728405 1.062129
1.506241 1.269373 1.345321 1.828694 1.029036 1.029036 0.721514
1.731853 1.406414 0.869982 0.901525 1.334125 1.334125 1.304915
0.5 0.772326 1.159405 1.773793 1.854496 0.691270 7.582796 1.691677
2.333372 1.381033 1.693753 1.954086 0.719767 10.713740 1.489551
0.906278 2.128215 2.517919 1.878350 0.807654 12.751710 1.440584
1.0 4.021173 1.505571 1.616389 4.601057 0.637286 1.205228 1.654745
3.552205 1.141802 0.760429 1.714344 0.204097 0.749852 0.648161
3.153475 1.869340 1.330379 3.146229 0.274938 0.863148 0.816919
3.0 1.730386 1.712858 2.823156 1.944915 1.290353 1.278683 0.441802
1.743547 0.958481 1.360395 0.609258 0.556973 0.900653 0.231205
1.531246 1.217893 1.133152 1.566374 1.034183 2.563201 0.544648
6.0 7.491243 4.959007 11.984900 14.824110 1.460042 14.339160 1.742513
8.709589 6.226643 11.173230 15.912830 1.610668 20.513880 2.250919
6.730411 5.592825 12.796570 15.055710 1.536903 20.236210 1.996716
9.0 1.550042 2.273255 0.611888 0.814029 0.874245 7.804387 1.216835
1.286587 2.985491 0.669221 0.808121 1.009184 5.403626 0.735552
1.763603 3.697727 0.740158 0.855448 0.901323 2.963197 1.151440
12.0 0.659497 1.284734 1.313399 0.988045 1.204775 8.536256 1.024598
3.250984 2.389733 1.484973 1.884897 1.850079 12.979700 1.120172
1.017229 0.672581 0.837188 0.465141 0.841363 6.292692 0.358020
冷胁迫
Cold stress		 0 0.383349 0.560141 0.854405 0.606571 0.728405 0.728405 1.062129
1.506241 1.269373 1.345321 1.828694 1.029036 1.029036 0.721514
1.731853 1.406414 0.869982 0.901525 1.334125 1.334125 1.304915
0.5 3.067984 0.073947 1.240688 0.905638 0.506053 3.463253 0.041140
2.764997 0.095788 1.552569 0.869109 0.680372 10.269000 0.048050
1.630744 0.025687 2.114081 0.600380 0.674204 4.344438 0.047003
1.0 6.912400 2.331709 2.603918 1.268249 1.842374 9.603889 0.064722
3.522932 4.848282 1.906822 0.534672 1.078122 4.360394 0.037754
1.921253 2.452198 0.514955 0.786965 1.011729 5.216602 0.027871
3.0 4.384372 1.205219 0.984415 1.079580 1.133988 7.848745 0.045085
4.254747 1.864312 1.894010 1.485352 1.642780 9.145513 0.052989
4.014151 2.783633 1.027342 0.985608 1.142598 12.440090 0.072667
6.0 2.086202 1.709914 1.716748 2.497598 1.202176 3.998536 0.074053
1.942518 1.154572 1.937527 2.730446 0.733247 4.476529 0.086771
1.385889 0.834515 1.365975 0.948892 0.348817 3.680676 0.047853
9.0 0.711963 0.238596 0.367345 1.016697 0.872451 1.058060 0.026753
1.105034 0.868088 0.321311 0.587120 1.104628 2.405538 0.033237
1.466575 1.280835 1.849609 1.070318 1.585804 5.327724 0.074461
12.0 5.369742 1.663197 2.324701 2.033130 0.741304 6.911496 0.062350
5.109689 0.758459 2.143919 2.832090 0.686679 5.455784 0.045538
4.807878 1.727933 3.264301 2.993412 0.647229 5.499532 0.046762
热胁迫
Heat stress		 0 0.383349 0.560141 0.854405 0.606571 0.728405 0.728405 1.062129
1.506241 1.269373 1.345321 1.828694 1.029036 1.029036 0.721514
1.731853 1.406414 0.869982 0.901525 1.334125 1.334125 1.304915
0.5 5.698402 0.342630 2.938812 1.912740 3.123081 0.395001 0.030419
6.523212 0.543866 4.800879 3.507437 3.619378 1.669364 0.089216
7.348022 0.435744 1.879438 2.008440 2.626784 0.813130 0.051139
1.0 11.545170 0.967400 0.065129 0.952527 3.284985 0.017801 0.001726
12.091210 0.729950 0.278504 1.931629 2.470521 0.042110 0.002321
12.637250 0.965469 0.173085 2.069349 4.086476 0.058591 0.004358
3.0 3.355991 0.049800 0.829518 3.078433 1.326899 0.576277 0.016437
8.287571 0.134510 3.354101 4.393673 1.842339 1.273716 0.029428
5.315243 0.866220 2.655746 5.708913 1.601412 0.721393 0.018568
6.0 0.043172 0.310586 0.386811 1.060621 2.231053 0.284953 0.006082
0.417197 1.199896 1.531503 2.426767 2.021476 0.704766 0.011086
0.364518 0.590544 1.353164 2.264082 1.811899 0.528078 0.007894
9.0 1.266708 0.243170 0.605378 3.033535 1.584369 0.668059 0.011319
3.190623 0.649720 1.499023 4.901491 1.326435 0.359377 0.018200
0.773340 0.517530 2.008419 3.327380 1.068478 0.485347 0.011061
12.0 0.300956 0.103220 0.158249 1.211129 0.470379 0.254347 0.002693
0.541412 0.579653 1.105782 1.806751 0.616462 0.383229 0.004141
0.320881 0.163824 0.260170 1.228762 0.557374 0.220774 0.003609
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