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作物学报

作物学报 ›› 2024, Vol. 50 ›› Issue (4): 897-813.doi: 10.3724/SP.J.1006.2024.31046

所属专题： 小麦：遗传育种·种质资源·分子遗传学

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

小麦及其祖先物种GRF转录因子家族鉴定与表达分析

王添宁1(), 冯雅岚2, 琚吉浩1, 吴毅1, 张均1, 马超1,*()   

  1. 1河南科技大学农学院, 河南洛阳 471000
    2武昌理工学院生命科学学院, 湖北武汉 430000
  • 收稿日期:2023-08-04 接受日期:2023-10-23 出版日期:2024-04-12 网络出版日期:2023-11-10
  • 通讯作者: * 马超, E-mail: machao840508@163.com
  • 作者简介:E-mail: haust6626wtn@163.com
  • 基金资助:
    国家自然科学基金项目(32372227);河南省自然科学基金项目(222300420430);河南省高等学校青年骨干教师培养计划(2021GGJS050)

Whole genome identification and analysis of GRFs transcription factor family in wheat and its ancestral species

WANG Tian-Ning1(), FENG Ya-Lan2, JU Ji-Hao1, WU Yi1, ZHANG Jun1, MA Chao1,*()   

  1. 1Henan University of Science and Technology, Agronomy College, Luoyang 471000, Henan, China
    2Wuchang University of Technology, College of Life Science, Wuhan 430223, Hubei, China
  • Received:2023-08-04 Accepted:2023-10-23 Published:2024-04-12 Published online:2023-11-10
  • Contact: * E-mail: machao840508@163.com
  • Supported by:
    National Natural Science Foundation of China(32372227);National Natural Science Foundation of Henan Province(222300420430);Training Program for University Young Key Teachers in Henan Province(2021GGJS050)

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摘要：

生长调控因子(growth-regulating factor, GRF)在植物的生长发育、逆境响应和激素信号转导中起着重要的调控作用。系统分析小麦及其祖先物种GRF转录因子家族成员在基因组上的分布、结构、进化以及表达特性, 对于深入研究GRF家族的生物学功能和小麦的进化具有重要的意义。本研究利用生物信息学方法, 对乌拉尔图小麦、拟斯卑尔脱山羊草、粗山羊草、栽培二粒小麦和普通小麦5个物种的GRF成员进行全基因组鉴定, 并对其理化性质、系统发育关系、基因结构、启动子顺式作用元件以及表达特性进行了分析。结果表明, 乌拉尔图小麦、拟斯卑尔脱山羊草、粗山羊草、栽培二粒小麦和普通小麦中分别有15、12、19、29和53个GRF成员, 通过种间共线性分析发现, TtGRFs分别有18个和29个成员与TuGRFs和AesGRFs具有共线性, TaGRFs分别有36个和37个成员与TtGRFs和AetGRFs具有共线性。启动子顺式作用元件预测发现GRF基因具有基本的转录元件以及一些与生长发育和逆境响应的结合元件。RT-qPCR分析表明, 多数GRF基因在外源IAA、GA和干旱胁迫条件下呈上调表达趋势, 而在高温胁迫条件下呈下调表达趋势, 表明GRF家族成员在响应激素和逆境胁迫中有重要作用。系统进化分析表明, 小麦与其祖先物种之间的GRF成员存在保守且复杂的进化关系。上述结果为GRF转录因子家族的进化及其功能研究提供了理论基础。

关键词: 小麦, 祖先物种, GRF, 生物信息学, 进化分析

Abstract:

Growth-regulating factors (GRFs) play important roles in plant growth, stress response, and hormone signal transduction. Systematic analysis of the distribution, structure, evolution, and expression characteristics of the GRF transcription factor family members in the genome of wheat and its ancestral species is of great significance for in-depth research on the biological functions of GRF family and the evolution of wheat. In this study, bioinformatics methods were used to identify the whole genome of GRF members from five species (Triticum urartu, Aegilops speltoides, Aegilops tauschii, Triticum turgidum, and Triticum aestivum), and their physical and chemical properties, phylogenetic relationships, gene structure, promoter cis-regulatory element, and expression characteristics were also analyzed. The results showed that there were 15, 12, 19, 29, and 53 GRF members in Triticum urartu, Aegilops speltoides, Aegilops tauschii, Triticum turgidum, and Triticum aestivum, respectively. Through interspecific colinearity analysis, we found that 18 and 29 members of TtGRFs were colinear with TuGRFs and AesGRFs, and 36 and 37 members of TaGRFs were colinear with TtGRFs and AetGRFs, respectively. The prediction of promoter cis-regulatory element found that GRF gene had basic transcription elements and some binding elements with growth, development, and stress response. RT-qPCR analysis revealed that most GRF genes up-regulated under exogenous IAA, GA, and drought stress, but down-regulated under high temperature stress, indicating that members of the GRF family exerted a crucial influence in hormone response and stress. Phylogenetic analysis evidenced that there was a conserved and complex evolutionary relationship between the GRF members of wheat and its ancestral species. The above results provide a theoretical basis for the evolution and functional research of the GRF transcription factor family.

Key words: wheat, ancestral species, GRF, bioinformatics, phylogenetic analysis

表1

本试验所用引物"

基因Gene 上游引物Forward primer (5'-3') 下游引物Reverse primer (5'-3')
TaGRF5-4A (XM_044510474.1) GAACCACCTTCTCCTTCGCCAG CCACAGGCTTTCTTGAACGG
TaGRF5-7A (XM_044569525.1) GCCTCGTCATACGGCAACAAAT AGAAGAAGCTCAGTTCACCCACG
TaGRF5-7D (XM_044584291.1) CAGGAAACAACAACAGGGGA CGGCTCAAAGTGGGAATACGA
TuGRF5-UN (XM_048693163.1) GCCTCGTCATACGGCAACAAAT AGAAGAAGCTCAGTTCACCCACG
AesGRF5-UN (AespeY2032UN01G008400.1) GAACCACCTTCTCCTTCGCCA CCACAGGCTTTCTTGAACGG
AetGRF5-7D (XM_020303029.3) CAGGAAACAACAACAGGGGA CGGCTCAAAGTGGGAATACGA
TtGRF5-4A (TRITD_4Av1G251150-2) GAACCACCTTCTCCTTCGCCAG CCACAGGCTTTCTTGAACGG
TtGRF5-7A (TRITD_7Av1G011750-2) GCCTCGTCATACGGCAACAAAT AGAAGAAGCTCAGTTCACCCACG
TaGRF11-2A (XM_044600790.1) ACACTCTGCTTGGACTTCGG CCACGATGCATATGGCGTTC
TaGRF11-2B (XM_044467676.1) TGGAAGAGTGTTACCGGTGC GGTTCTGGGCTCTTCCCAAA
TaGRF11-2D (XM_044476137.1) CTGAAGGCGCGCAGAATTAC TCTCCAGCATCGCCACTTTT
TuGRF11-2A (XM_048703523.1) ACACTCTGCTTGGACTTCGG CCACGATGCATATGGCGTTC
AesGRF11-2B (AespeY2032CH2S01G240700.1) TGGAAGAGTGTTACCGGTGC GGTTCTGGGCTCTTCCCAAA
AetGRF11-2D (XM_020298039.3) CTGAAGGCGCGCAGAATTAC TCTCCAGCATCGCCACTTTT
TtGRF11-2A (TRITD_2Av1G117900) ACACTCTGCTTGGACTTCGG CCACGATGCATATGGCGTTC
TtGRF11-2B (TRITD_2Bv1G105550-2) TGGAAGAGTGTTACCGGTGC GGTTCTGGGCTCTTCCCAAA
Actin (XM_044554036.1) TATGCCAGCGGTCGAACAAC GGAACAGCACCTCAGGGCAC
GAPDH (AK456344.1) TAAGGGTGGTGCCAAGAAGGT AGCAAGAGGAGCAAGGCAGTT

附表1

小麦及其祖先物种GRF蛋白理化性质和亚细胞定位"

基因名称
Gene name		 登录号
Accession number		 氨基酸数目
Number of amino acid		 相对分子质量
Molecular weight		 等电点
Theoretical pI		 不稳定指数
Instability index		 脂肪系数
Aliphatic index		 平均疏水指数
Grand average of hydropathicity		 亚细胞定位
Subcellular localization
TuGRF10-6A (XM_048677422.1) XM_048677422.1 212 22605.56 9.64 57.97 67.83 −0.393 nucleus
TuGRF13-6A (XM_048677685.1) XM_048677685.1 309 32548.58 10.01 59.63 66.34 −0.760 chloroplast
TuGRF1-6A (XM_048677760.1) XM_048677760.1 409 44786.45 7.22 51.42 46.65 −0.833 nucleus
TuGRF2-7A (XM_048685991.1) XM_048685991.1 311 34358.18 8.55 65.88 51.03 −0.841 nucleus
TuGRF2-7A (XM_048685992.1) XM_048685992.1 310 34287.11 8.55 66.34 50.87 −0.850 nucleus
TuGRF5-UN (XM_048693163.1) XM_048693163.1 370 40016.38 8.78 60.44 46.24 −0.760 nucleus
TuGRF14-UN (XM_048693245.1) XM_048693245.1 290 31954.68 7.57 59.98 54.21 −0.899 nucleus
TuGRF6-UN (XM_048693253.1) XM_048693253.1 607 63999.27 6.72 51.77 64.93 −0.428 nucleus
TuGRF8-UN (XM_048695887.1) XM_048695887.1 196 21264.79 8.59 39.77 63.47 −0.640 chloroplast
TuGRF12-2A (XM_048697161.1) XM_048697161.1 225 23851.86 9.68 51.60 64.36 −0.448 nucleus
TuGRF3-2A (XM_048703059.1) XM_048703059.1 384 42335.86 6.78 60.09 54.35 −0.628 nucleus
TuGRF11-2A (XM_048703523.1) XM_048703523.1 265 28286.31 4.72 55.17 62.26 −0.668 nucleus
TuGRF9-4A (XM_048713185.1) XM_048713185.1 408 45353.73 9.00 62.33 55.54 −0.850 nucleus
TuGRF15-4A (XM_048716712.1) XM_048716712.1 395 43318.03 6.99 68.77 55.54 −0.787 nucleus
TuGRF15-4A (XM_048718295.1) XM_048718295.1 390 42618.24 7.90 61.82 65.08 −0.653 mitochondrion
AesGRF2-7B (AespeY2032CH7S01G162000.1) AespeY2032CH7S01G162000.1 310 34313.19 9.05 64.92 50.55 −0.851 nucleus
AesGRF2-7B (AespeY2032CH7S01G147800.1) AespeY2032CH7S01G147800.1 315 34849.78 9.05 62.70 51.30 −0.841 nucleus
AesGRF6-6B (AespeY2032CH6S01G269500.1) AespeY2032CH6S01G269500.1 311 33561.65 8.12 40.72 67.97 −0.412 mitochondrion
AesGRF1-6B (AespeY2032CH6S01G467300.1) AespeY2032CH6S01G467300.1 307 33626.73 9.52 66.07 46.22 −0.735 nucleus
AesGRF9-4B (AespeY2032CH4S01G027900.1) AespeY2032CH4S01G027900.1 403 44535.84 9.01 61.60 56.03 −0.818 nucleus
AesGRF6-4B (AespeY2032CH4S01G078800.1) AespeY2032CH4S01G078800.1 615 64726.95 6.74 53.11 64.42 −0.433 nucleus
AesGRF3-2B (AespeY2032CH2S01G448800.1) AespeY2032CH2S01G448800.1 356 39440.66 7.00 54.62 53.90 −0.692 nucleus
AesGRF12-2B (AespeY2032CH2S01G398500.1) AespeY2032CH2S01G398500.1 229 24236.21 9.54 49.84 63.23 −0.470 nucleus
AesGRF11-2B (AespeY2032CH2S01G240700.1) AespeY2032CH2S01G240700.1 263 28126.14 4.72 55.31 63.12 −0.685 nucleus
AesGRF5-UN (AespeY2032UN01G008400.1) AespeY2032UN01G008400.1 266 28604.02 9.14 56.79 56.32 −0.624 nucleus
AesGRF14-1B (AespeY2032CH1S01G408000.1) AespeY2032CH1S01G408000.1 360 38840.69 8.28 66.92 56.17 −0.784 chloroplast
AesGRF10-6B (AespeY2032CH6S01G376400.1) AespeY2032CH6S01G376400.1 176 18162.52 10.83 61.40 70.23 −0.286 nucleus
AetGRF2-7D (XM_020291396.3) XM_020291396.3 321 35476.32 8.26 63.25 50.65 −0.873 nucleus
AetGRF2-7D (XM_020291397.3) XM_020291397.3 320 35405.24 8.26 63.68 50.50 −0.882 nucleus
AetGRF11-2D (XM_020298039.3) XM_020298039.3 264 28168.18 4.75 55.20 62.16 −0.692 nucleus
AetGRF1-6D (XM_020300487.3) XM_020300487.3 414 45263.95 7.24 51.94 46.09 −0.835 nucleus
AetGRF5-7D (XM_020303029.3) XM_020303029.3 366 39566.92 8.78 57.38 47.02 −0.727 nucleus
AetGRF3-2D (XM_020308591.3) XM_020308591.3 393 42893.44 7.04 62.06 52.88 −0.621 nucleus
AetGRF4-6D (XM_020309598.3) XM_020309598.3 409 43620.48 8.16 64.18 52.40 −0.559 nucleus
AetGRF6-6D (XM_020319993.2) XM_020319993.2 311 33231.24 6.85 44.20 66.78 −0.402 mitochondrion
AetGRF10-6D (XM_020324838.3) XM_020324838.3 215 22773.80 9.90 55.13 68.74 −0.359 nucleus
AetGRF6-4D (XM_020331699.3) XM_020331699.3 578 61162.10 6.58 53.81 64.43 −0.450 nucleus
AetGRF12-2D (XM_020332526.3) XM_020332526.3 229 24221.30 9.57 49.91 64.10 −0.443 nucleus
AetGRF5-7D (XM_040395388.2) XM_040395388.2 330 36154.22 8.58 54.38 46.24 −0.754 nucleus
AetGRF13-6D (XM_020297920.3) XM_020297920.3 304 32077.13 9.79 60.82 63.95 −0.754 nucleus
AetGRF13-7D (XM_020306383.3) XM_020306383.3 255 26078.93 10.18 54.79 55.33 −0.763 chloroplast
AetGRF13-7D (XM_020306384.3) XM_020306384.3 254 25991.86 10.18 54.21 55.55 −0.763 chloroplast
AetGRF15-5D (XM_020330238.3) XM_020330238.3 389 42721.28 6.99 63.51 55.94 −0.769 mitochondrion
AetGRF15-5D (XM_020330240.3) XM_020330240.3 404 44491.00 6.17 64.90 56.04 −0.821 nucleus
AetGRF14-1D (XM_040391509.2) XM_040391509.2 361 39248.11 7.56 71.15 56.84 −0.761 nucleus
AetGRF15-5D (XM_045229328.1) XM_045229328.1 397 43646.97 6.16 66.28 54.58 −0.846 nucleus
TtGRF14-1A (TRITD_1Av1G209470) TRITD_1Av1G209470 365 39597.35 8.00 67.98 54.60 −0.823 nucleus
TtGRF14-1B (TRITD_1Bv1G214800) TRITD_1Bv1G214800 339 36993.59 8.01 60.06 59.32 −0.742 nucleus
TtGRF13-2A (TRITD_2Av1G077710-2) TRITD_2Av1G077710-2 627 67054.64 4.85 68.21 55.06 −0.983 nucleus
TtGRF11-2A (TRITD_2Av1G117900) TRITD_2Av1G117900 260 27872.84 4.72 56.04 61.19 −0.715 nucleus
TtGRF12-2A (TRITD_2Av1G235170) TRITD_2Av1G235170 225 23793.83 9.82 51.22 64.36 −0.434 nucleus
TtGRF3-2A (TRITD_2Av1G251570) TRITD_2Av1G251570 384 42326.85 6.76 61.22 54.61 −0.623 nucleus
TtGRF11-2B (TRITD_2Bv1G105550-2) TRITD_2Bv1G105550-2 263 28160.15 4.72 54.07 61.63 −0.689 nucleus
TtGRF12-2B (TRITD_2Bv1G197140) TRITD_2Bv1G197140 227 24103.25 9.82 51.75 64.23 −0.459 chloroplast
TtGRF3-2B (TRITD_2Bv1G214250) TRITD_2Bv1G214250 387 42478.01 7.03 61.20 52.40 −0.620 nucleus
TtGRF6-4A (TRITD_4Av1G187110-2) TRITD_4Av1G187110-2 607 63953.28 6.87 51.64 65.58 −0.416 nucleus
TtGRF9-4A (TRITD_4Av1G200390-2) TRITD_4Av1G200390-2 433 48568.80 9.19 57.27 64.27 −0.661 nucleus
TtGRF15-4A (TRITD_4Av1G222610) TRITD_4Av1G222610 393 43127.45 5.51 67.44 56.13 −0.818 nucleus
TtGRF15-4A (TRITD_4Av1G223320) TRITD_4Av1G223320 391 42951.53 6.99 68.24 54.37 −0.794 nucleus
TtGRF5-4A (TRITD_4Av1G251150-2) TRITD_4Av1G251150-2 371 40019.38 8.77 62.57 46.39 −0.748 nucleus
TtGRF9-4B (TRITD_4Bv1G006160) TRITD_4Bv1G006160 404 44678.98 8.93 60.76 56.58 −0.799 nucleus
TtGRF6-4B (TRITD_4Bv1G019740-2) TRITD_4Bv1G019740-2 611 64277.53 6.72 51.31 65.01 −0.423 nucleus
TtGRF10-6A (TRITD_6Av1G164980) TRITD_6Av1G164980 212 22625.61 9.54 52.82 68.25 −0.373 nucleus
TtGRF4-6A (TRITD_6Av1G170910) TRITD_6Av1G170910 408 43471.22 7.65 63.22 52.28 −0.573 nucleus
TtGRF13-6A (TRITD_6Av1G195840) TRITD_6Av1G195840 184 19379.22 10.66 45.82 77.39 −0.454 cytosol
TtGRF1-6A (TRITD_6Av1G201020-3) TRITD_6Av1G201020-3 409 44786.45 7.22 51.42 46.65 −0.833 nucleus
TtGRF10-6B (TRITD_6Bv1G150350) TRITD_6Bv1G150350 211 22347.22 9.64 53.20 67.68 −0.382 nucleus
TtGRF4-6B (TRITD_6Bv1G157700-2) TRITD_6Bv1G157700-2 408 43678.55 8.66 65.27 52.77 −0.581 nucleus
TtGRF1-6B (TRITD_6Bv1G196330-3) TRITD_6Bv1G196330-3 410 44724.40 7.21 51.02 45.83 −0.821 nucleus
TtGRF5-7A (TRITD_7Av1G011750-2) TRITD_7Av1G011750-2 279 30182.21 8.83 58.80 44.09 −0.865 nucleus
TtGRF2-7A (TRITD_7Av1G053020-2) TRITD_7Av1G053020-2 310 34199.09 8.93 65.08 51.48 −0.813 nucleus
TtGRF2-7B (TRITD_7Bv1G027670) TRITD_7Bv1G027670 316 34886.68 8.55 66.25 49.59 −0.882 nucleus
TtGRF6-4A (TRITD_4Av1G187110) TRITD_4Av1G187110 490 52799.36 6.16 51.77 73.71 −0.274 mitochondrion
TtGRF3-6A (TRITD_6Av1G013860) TRITD_6Av1G013860 247 27740.32 5.00 77.10 79.80 −0.456 nucleus
TtGRF3-6A (TRITD_6Av1G211060) TRITD_6Av1G211060 247 28061.02 5.07 70.80 87.25 −0.388 nucleus
TaGRF12-2B (XM_044464321.1) XM_044464321.1 227 24077.17 9.82 53.68 62.51 −0.479 nucleus
TaGRF11-2B (XM_044467676.1) XM_044467676.1 263 28160.15 4.72 54.07 61.63 −0.689 nucleus
TaGRF11-2B (XM_044467677.1) XM_044467677.1 258 27746.68 4.72 54.92 60.54 −0.736 nucleus
TaGRF3-2B (XM_044469953.1) XM_044469953.1 387 42454.97 7.01 61.92 52.40 −0.621 nucleus
TaGRF12-2D (XM_044472604.1) XM_044472604.1 229 24221.30 9.57 49.91 64.10 −0.443 nucleus
TaGRF11-2D (XM_044476137.1) XM_044476137.1 264 28182.20 4.76 55.93 62.16 −0.692 nucleus
TaGRF3-2D (XM_044478186.1) XM_044478186.1 391 42780.33 7.04 60.85 53.40 −0.620 nucleus
TaGRF14-1A (XM_044491129.1) XM_044491129.1 364 39469.22 8.00 67.24 54.75 −0.816 nucleus
TaGRF15-4A (XM_044505037.1) XM_044505037.1 391 42973.46 6.63 68.13 55.12 −0.805 nucleus
TaGRF9-4A (XM_044505809.1) XM_044505809.1 546 60568.73 9.15 68.70 53.13 −0.916 nucleus
TaGRF6-4A (XM_044508878.1) XM_044508878.1 607 63953.28 6.87 51.64 65.58 −0.416 nucleus
TaGRF15-4A (XM_044509907.1) XM_044509907.1 391 42920.51 6.99 68.10 54.37 −0.789 nucleus
TaGRF5-4A (XM_044510474.1) XM_044510474.1 371 39941.33 8.50 61.23 47.44 −0.726 nucleus
TaGRF9-4B (XM_044511958.1) XM_044511958.1 404 44678.98 8.93 60.76 56.58 −0.799 nucleus
TaGRF6-4B (XM_044512340.1) XM_044512340.1 611 64277.53 6.72 51.31 65.01 −0.423 nucleus
TaGRF6-4B (XM_044512341.1) XM_044512341.1 598 62948.00 6.60 52.00 64.13 −0.442 nucleus
TaGRF6-4B (XM_044512342.1) XM_044512342.1 579 60297.62 9.29 56.44 69.46 −0.442 nucleus
TaGRF6-4D (XM_044517570.1) XM_044517570.1 578 61162.10 6.58 53.81 64.43 −0.450 nucleus
TaGRF15-5B (XM_044538321.1) XM_044538321.1 377 42148.30 9.60 68.01 59.55 −0.707 nucleus
TaGRF13-5D (XM_044540830.1) XM_044540830.1 199 22033.98 4.25 72.48 58.84 −0.863 cytosol
TaGRF15-5D (XM_044544157.1) XM_044544157.1 404 44491.00 6.17 64.90 56.04 −0.821 nucleus
TaGRF15-5D (XM_044546482.1) XM_044546482.1 389 42721.28 6.99 63.51 55.94 −0.769 mitochondrion
TaGRF4-6A (XM_044548020.1) XM_044548020.1 408 43457.29 7.65 65.49 52.03 −0.579 nucleus
TaGRF1-6A (XM_044548647.1) XM_044548647.1 409 44786.45 7.22 51.42 46.65 −0.833 nucleus
TaGRF10-6A (XM_044550602.1) XM_044550602.1 212 22597.56 9.54 52.41 67.36 −0.384 nucleus
TaGRF13-6A (XM_044550808.1) XM_044550808.1 304 31967.89 10.50 58.17 64.87 −0.784 chloroplast
TaGRF10-6B (XM_044553553.1) XM_044553553.1 211 22347.22 9.64 53.20 67.68 −0.382 nucleus
TaGRF13-6B (XM_044553828.1) XM_044553828.1 318 33481.63 9.57 64.42 63.90 −0.753 nucleus
TaGRF4-6B (XM_044557754.1) XM_044557754.1 406 43435.28 8.46 64.46 53.03 −0.571 nucleus
TaGRF1-6B (XM_044558455.1) XM_044558455.1 410 44724.40 7.21 51.02 45.83 −0.821 nucleus
TaGRF6-6D (XM_044560555.1) XM_044560555.1 311 33231.24 6.85 44.20 66.78 −0.402 mitochondrion
TaGRF10-6D (XM_044560736.1) XM_044560736.1 215 22750.77 9.90 55.13 68.74 −0.360 nucleus
TaGRF4-6D (XM_044564540.1) XM_044564540.1 409 43620.48 8.16 64.18 52.40 −0.559 nucleus
TaGRF13-6D (XM_044565096.1) XM_044565096.1 304 32105.19 9.79 60.82 64.57 −0.746 nucleus
TaGRF1-6D (XM_044565199.1) XM_044565199.1 414 45263.95 7.24 51.94 46.09 −0.835 nucleus
TaGRF14-1B (XM_044565877.1) XM_044565877.1 364 39490.49 7.55 67.53 56.87 −0.775 nucleus
TaGRF5-7A (XM_044569525.1) XM_044569525.1 370 40161.60 8.78 59.57 46.24 −0.762 nucleus
TaGRF2-7A (XM_044571651.1) XM_044571651.1 310 34285.13 8.55 65.17 51.48 −0.832 nucleus
TaGRF2-7A (XM_044571652.1) XM_044571652.1 309 34214.05 8.55 65.62 51.33 −0.841 nucleus
TaGRF2-7B (XM_044575454.1) XM_044575454.1 316 34886.68 8.55 66.25 49.59 −0.882 nucleus
TaGRF13-7D (XM_044583768.1) XM_044583768.1 255 26078.93 10.18 54.79 55.33 −0.763 chloroplast
TaGRF13-7D (XM_044583769.1) XM_044583769.1 254 25991.86 10.18 54.21 55.55 −0.763 chloroplast
TaGRF5-7D (XM_044584291.1) XM_044584291.1 368 39895.24 8.57 58.69 46.77 −0.750 nucleus
TaGRF2-7D (XM_044587055.1) XM_044587055.1 321 35476.32 8.26 63.25 50.65 −0.873 nucleus
TaGRF2-7D (XM_044587056.1) XM_044587056.1 320 35405.24 8.26 63.68 50.50 −0.882 nucleus
TaGRF14-1D (XM_044595886.1) XM_044595886.1 361 39205.09 7.04 69.57 57.92 −0.739 chloroplast
TaGRF14-1D (XM_044595887.1) XM_044595887.1 361 39205.09 7.04 69.57 57.92 −0.739 chloroplast
TaGRF14-1D (XM_044595888.1) XM_044595888.1 361 39205.09 7.04 69.57 57.92 −0.739 chloroplast
TaGRF12-2A (XM_044597639.1) XM_044597639.1 225 23823.85 9.82 54.41 64.36 −0.436 nucleus
TaGRF11-2A (XM_044600790.1) XM_044600790.1 265 28286.31 4.72 55.17 62.26 −0.668 nucleus
TaGRF3-2A (XM_044602884.1) XM_044602884.1 384 42326.85 6.76 61.22 54.61 −0.623 nucleus
TaGRF3-6A (XM_044551538.1) XM_044551538.1 108 12319.67 5.62 73.33 66.94 −0.825 mitochondrion
TaGRF3-6A (XM_044551539.1) XM_044551539.1 107 12191.54 5.62 72.12 67.57 −0.800 mitochondrion

附表2

小麦及其祖先物种GRF的二级结构预测"

基因名称
Gene Name		 α-螺旋
Alpha helix (Hh)		 β-转角
Beta turn (Tt)		 延伸链
Extended strand (Ee)		 无规则卷曲
Random coil (Cc)
TuGRF10-6A (XM_048677422.1) 24.53 4.25 13.21 58.02
TuGRF13-6A (XM_048677685.1) 16.50 8.09 12.30 63.11
TuGRF1-6A (XM_048677760.1) 16.63 5.62 9.78 67.97
TuGRF2-7A (XM_048685991.1) 27.33 4.18 7.72 60.77
TuGRF2-7A (XM_048685992.1) 26.45 4.84 8.06 60.65
TuGRF5-UN (XM_048693163.1) 20.27 4.05 9.46 66.22
TuGRF14-UN (XM_048693245.1) 31.38 3.10 9.66 55.86
TuGRF6-UN (XM_048693253.1) 26.03 3.13 8.07 62.77
TuGRF8-UN (XM_048695887.1) 43.37 9.18 10.71 36.73
TuGRF12-2A (XM_048697161.1) 20.44 4.44 9.33 65.78
TuGRF3-2A (XM_048703059.1) 23.70 3.12 5.73 67.45
TuGRF11-2A (XM_048703523.1) 35.85 7.17 13.58 43.40
TuGRF9-4A (XM_048713185.1) 25.25 2.45 8.33 63.97
TuGRF15-4A (XM_048716712.1) 25.06 0.51 6.58 67.85
TuGRF15-4A (XM_048718295.1) 25.13 2.56 8.97 63.33
AesGRF2-7B (AespeY2032CH7S01G162000.1) 27.42 4.84 7.74 60.00
AesGRF2-7B (AespeY2032CH7S01G147800.1) 26.35 4.44 7.94 61.27
AesGRF6-6B (AespeY2032CH6S01G269500.1) 37.94 6.75 14.47 40.84
AesGRF1-6B (AespeY2032CH6S01G467300.1) 16.29 7.82 13.68 62.21
AesGRF9-4B (AespeY2032CH4S01G027900.1) 26.80 4.47 8.19 60.55
AesGRF6-4B (AespeY2032CH4S01G078800.1) 25.85 4.23 11.54 58.37
AesGRF3-2B (AespeY2032CH2S01G448800.1) 23.88 3.37 7.58 65.17
AesGRF12-2B (AespeY2032CH2S01G398500.1) 20.09 6.11 12.23 61.57
AesGRF11-2B (AespeY2032CH2S01G240700.1) 31.56 6.08 14.07 48.29
AesGRF5-UN (AespeY2032UN01G008400.1) 16.54 4.51 6.02 72.93
AesGRF14-1B (AespeY2032CH1S01G408000.1) 29.44 2.22 7.78 60.56
AesGRF10-6B (AespeY2032CH6S01G376400.1) 35.80 4.55 11.36 48.30
AetGRF2-7D (XM_020291396.3) 26.48 3.74 6.54 63.24
AetGRF2-7D (XM_020291397.3) 24.06 2.81 7.81 65.31
AetGRF11-2D (XM_020298039.3) 34.09 7.95 15.91 42.05
AetGRF1-6D (XM_020300487.3) 20.29 5.31 10.14 64.25
AetGRF5-7D (XM_020303029.3) 15.30 3.83 7.92 72.95
AetGRF3-2D (XM_020308591.3) 22.90 3.82 5.09 68.19
AetGRF4-6D (XM_020309598.3) 18.83 3.91 5.38 71.88
AetGRF6-6D (XM_020319993.2) 42.12 8.36 13.50 36.01
AetGRF10-6D (XM_020324838.3) 32.09 6.05 11.63 50.23
AetGRF6-4D (XM_020331699.3) 25.26 3.81 9.34 61.59
AetGRF12-2D (XM_020332526.3) 24.02 6.11 9.61 60.26
AetGRF5-7D (XM_040395388.2) 15.76 3.94 9.39 70.91
AetGRF13-6D (XM_020297920.3) 14.80 8.55 13.16 63.49
AetGRF13-7D (XM_020306383.3) 25.49 5.49 11.76 57.25
AetGRF13-7D (XM_020306384.3) 20.87 6.69 10.24 62.20
AetGRF15-5D (XM_020330238.3) 31.11 2.31 8.48 58.10
AetGRF15-5D (XM_020330240.3) 26.73 1.49 9.65 62.13
AetGRF14-1D (XM_040391509.2) 29.92 1.39 6.65 62.05
AetGRF15-5D (XM_045229328.1) 23.93 0.76 10.08 65.24
TtGRF14-1A (TRITD_1Av1G209470) 30.41 3.01 7.40 59.18
TtGRF14-1B (TRITD_1Bv1G214800) 31.13 2.48 8.26 58.13
TtGRF13-2A (TRITD_2Av1G077710-2) 33.49 6.54 9.25 50.72
TtGRF11-2A (TRITD_2Av1G117900) 31.92 7.31 15.77 45.00
TtGRF12-2A (TRITD_2Av1G235170) 20.00 4.44 9.33 66.22
TtGRF3-2A (TRITD_2Av1G251570) 27.08 3.12 4.95 64.84
TtGRF11-2B (TRITD_2Bv1G105550-2) 33.46 6.84 13.31 46.39
TtGRF12-2B (TRITD_2Bv1G197140) 18.94 5.29 8.37 67.40
TtGRF3-2B (TRITD_2Bv1G214250) 21.45 3.36 5.17 70.03
TtGRF6-4A (TRITD_4Av1G187110-2) 27.51 3.62 8.90 59.97
TtGRF9-4A (TRITD_4Av1G200390-2) 28.18 5.08 10.16 56.58
TtGRF15-4A (TRITD_4Av1G222610) 23.41 2.04 8.65 65.90
TtGRF15-4A (TRITD_4Av1G223320) 26.85 1.28 7.16 64.71
TtGRF5-4A (TRITD_4Av1G251150-2) 19.95 4.04 11.05 64.96
TtGRF9-4B (TRITD_4Bv1G006160) 22.77 2.72 9.41 65.10
TtGRF6-4B (TRITD_4Bv1G019740-2) 23.90 4.42 10.31 61.37
TtGRF10-6A (TRITD_6Av1G164980) 29.25 2.83 11.32 56.60
TtGRF4-6A (TRITD_6Av1G170910) 19.61 2.45 6.62 71.32
TtGRF13-6A (TRITD_6Av1G195840) 16.85 10.87 16.85 55.43
TtGRF1-6A (TRITD_6Av1G201020-3) 16.63 5.62 9.78 67.97
TtGRF10-6B (TRITD_6Bv1G150350) 32.70 2.84 8.53 55.92
TtGRF4-6B (TRITD_6Bv1G157700-2) 14.46 3.19 7.60 74.75
TtGRF1-6B (TRITD_6Bv1G196330-3) 19.76 6.34 13.17 60.73
TtGRF5-7A (TRITD_7Av1G011750-2) 15.41 3.23 6.09 75.27
TtGRF2-7A (TRITD_7Av1G053020-2) 26.13 4.52 8.06 61.29
TtGRF2-7B (TRITD_7Bv1G027670) 27.85 3.80 6.96 61.39
TtGRF6-4A (TRITD_4Av1G187110) 27.96 5.92 17.14 48.98
TtGRF3-6A (TRITD_6Av1G013860) 38.46 5.67 8.50 47.37
TtGRF3-6A (TRITD_6Av1G211060) 36.44 4.86 10.53 48.18
TaGRF12-2B (XM_044464321.1) 18.94 5.29 9.69 66.08
TaGRF11-2B (XM_044467676.1) 33.46 6.84 13.31 46.39
TaGRF11-2B (XM_044467677.1) 32.56 7.75 12.79 46.90
TaGRF3-2B (XM_044469953.1) 24.29 4.39 8.27 63.05
TaGRF12-2D (XM_044472604.1) 24.02 6.11 9.61 60.26
TaGRF11-2D (XM_044476137.1) 34.09 7.95 15.91 42.05
TaGRF3-2D (XM_044478186.1) 23.02 3.58 5.37 68.03
TaGRF14-1A (XM_044491129.1) 29.40 3.02 7.69 59.89
TaGRF15-4A (XM_044505037.1) 26.09 1.79 7.93 64.19
TaGRF9-4A (XM_044505809.1) 24.36 4.03 10.26 61.36
TaGRF6-4A (XM_044508878.1) 27.51 3.62 8.90 59.97
TaGRF15-4A (XM_044509907.1) 26.09 0.77 8.18 64.96
TaGRF5-4A (XM_044510474.1) 16.17 4.04 10.51 69.27
TaGRF9-4B (XM_044511958.1) 22.77 2.72 9.41 65.10
TaGRF6-4B (XM_044512340.1) 23.90 4.42 10.31 61.37
TaGRF6-4B (XM_044512341.1) 24.75 3.18 10.54 61.54
TaGRF6-4B (XM_044512342.1) 37.48 4.84 10.02 47.67
TaGRF6-4D (XM_044517570.1) 25.26 3.81 9.34 61.59
TaGRF15-5B (XM_044538321.1) 22.55 3.45 7.96 66.05
TaGRF13-5D (XM_044540830.1) 30.65 4.02 16.08 49.25
TaGRF15-5D (XM_044544157.1) 26.73 1.49 9.65 62.13
TaGRF15-5D (XM_044546482.1) 31.11 2.31 8.48 58.10
TaGRF4-6A (XM_044548020.1) 16.18 3.43 7.35 73.04
TaGRF1-6A (XM_044548647.1) 16.63 5.62 9.78 67.97
TaGRF10-6A (XM_044550602.1) 26.42 4.25 14.62 54.72
TaGRF13-6A (XM_044550808.1) 16.45 8.22 12.83 62.50
TaGRF10-6B (XM_044553553.1) 32.70 2.84 8.53 55.92
TaGRF13-6B (XM_044553828.1) 16.98 5.97 10.06 66.98
TaGRF4-6B (XM_044557754.1) 17.49 2.96 7.88 71.67
TaGRF1-6B (XM_044558455.1) 19.76 6.34 13.17 60.73
TaGRF6-6D (XM_044560555.1) 42.12 8.36 13.50 36.01
TaGRF10-6D (XM_044560736.1) 29.77 5.58 12.56 52.09
TaGRF4-6D (XM_044564540.1) 18.83 3.91 5.38 71.88
TaGRF13-6D (XM_044565096.1) 14.80 8.55 13.16 63.49
TaGRF1-6D (XM_044565199.1) 20.29 5.31 10.14 64.25
TaGRF14-1B (XM_044565877.1) 29.40 1.65 8.24 60.71
TaGRF5-7A (XM_044569525.1) 24.05 3.78 8.38 63.78
TaGRF2-7A (XM_044571651.1) 26.13 4.84 6.77 62.26
TaGRF2-7A (XM_044571652.1) 26.86 3.24 7.44 62.46
TaGRF2-7B (XM_044575454.1) 27.85 3.80 6.96 61.39
TaGRF13-7D (XM_044583768.1) 25.49 5.49 11.76 57.25
TaGRF13-7D (XM_044583769.1) 20.87 6.69 10.24 62.20
TaGRF5-7D (XM_044584291.1) 18.21 11.96 11.96 66.30
TaGRF2-7D (XM_044587055.1) 26.48 3.74 6.54 63.24
TaGRF2-7D (XM_044587056.1) 24.06 2.81 7.81 65.31
TaGRF14-1D (XM_044595886.1) 29.92 2.22 7.48 60.39
TaGRF14-1D (XM_044595887.1) 29.92 2.22 7.48 60.39
TaGRF14-1D (XM_044595888.1) 29.92 2.22 7.48 60.39
TaGRF12-2A (XM_044597639.1) 32.44 6.22 9.78 51.56
TaGRF11-2A (XM_044600790.1) 35.85 7.17 13.58 43.40
TaGRF3-2A (XM_044602884.1) 27.08 3.12 4.95 64.84
TaGRF3-6A (XM_044551538.1) 38.89 11.11 4.63 45.37
TaGRF3-6A (XM_044551539.1) 33.64 11.21 7.48 47.66

图1

小麦及其祖先物种GRF5的二级结构和三级结构预测"

图2

小麦及其祖先物种GRF家族成员的系统进化树"

图3

小麦及其祖先物种GRF基因的染色体定位"

图4

小麦及其祖先物种GRF基因结构和保守结构域分析"

图5

小麦及其祖先物种GRF基因启动子顺式作用元件"

图6

小麦及其祖先物种GRF基因的染色体位置及共线性关系 所有共线性区域和基因由灰线连接, 红线表示GRF基因的片段重复基因对。"

图7

小麦及其祖先物种间GRF基因的共线性关系"

附表3

小麦及其祖先物种GRF基因Ka/Ks分析"

非同义替换率
Ka		 同义替换率
Ks		 非同义替换/同义替换
Ka/Ks		 有效长度
Effective length		 平均S位点
Average S-sites
0.1863 0.4703 0.3962 618 162.9167
0.3536 0.5820 0.6077 507 138.7500
0.0087 0.0343 0.2543 930 238.5000
0.3058 0.3947 0.7748 894 223.7500
0.2048 0.4338 0.4720 1101 277.5000
0.1760 0.4206 0.4183 612 163.5833
0.0441 0.1302 0.3385 1053 246.8333
0.0068 0.0450 0.1509 774 183.0000
0.0122 0.0950 0.1284 675 179.2500
0.1970 0.3967 0.4967 612 162.8333
0.1833 0.4039 0.4538 612 163.5833
0.0346 0.0753 0.4596 1137 279.2500
0.1968 0.4185 0.4704 612 162.9167
0.1812 0.3579 0.5062 603 161.3333
0.1962 0.4414 0.4445 1080 270.5000
0.1884 0.4276 0.4406 1074 270.2500
0.0595 0.2097 0.2839 1173 261.5000
0.0220 0.0312 0.7073 624 163.7500
0.0236 0.0869 0.2714 1209 304.6667
0.0043 0.0896 0.0484 1221 296.1667
0.3078 0.4399 0.6997 888 221.5833
0.3095 0.4099 0.7550 888 221.9167
0.0194 0.0441 0.4401 912 233.5000
0.0060 0.0818 0.0736 681 180.5833
0.1910 0.4081 0.4681 612 162.8333
0.1782 0.3482 0.5119 603 161.2500
0.1786 0.4224 0.4228 612 163.0833
0.0281 0.0546 0.5148 1155 284.9167
0.1916 0.4266 0.4492 1065 268.1667
0.1893 0.4372 0.4329 1074 270.4167
0.1897 0.4357 0.4354 1074 270.0833
0.1870 0.3705 0.5047 612 163.3333
0.1762 0.3775 0.4668 612 164.0833
0.1788 0.4100 0.4361 612 163.5833
0.2148 0.4374 0.4909 1107 278.3333
0.2136 0.4598 0.4647 1095 275.1667
0.2166 0.4457 0.4858 1101 276.5000
0.0414 0.1427 0.2904 1089 257.8333
0.0274 0.0738 0.3710 1074 256.0000
0.0156 0.1087 0.1439 1209 271.8333
0.0112 0.0971 0.1154 1800 450.3333
0.0116 0.0776 0.1497 1734 433.8333
0.0122 0.0978 0.1244 1101 272.5833
0.0109 0.0891 0.1226 1104 273.8333
0.0108 0.0777 0.1395 1734 433.3333
0.0237 0.0905 0.2615 1206 304.9167
0.0331 0.0735 0.4507 1218 307.2500
0.0043 0.0896 0.0484 1221 296.1667
0.0032 0.0821 0.0393 1227 295.8333
0.3071 0.3902 0.7872 894 224.0833
0.0198 0.0312 0.6356 624 163.7500
0.0216 0.0432 0.4998 636 166.5833
0.0648 0.1136 0.5705 888 227.6667
0.0593 0.1258 0.4715 873 224.4167
0.4300 0.6799 0.6324 678 176.3333
0.0195 0.0497 0.3931 633 166.3333
0.0303 0.0534 0.5680 906 232.9167
0.4408 0.6248 0.7055 672 176.5000
0.0134 0.0651 0.2063 1206 304.6667
0.0054 0.0739 0.0729 1230 298.3333
0.3095 0.4099 0.7550 888 221.9167
0.3124 0.3876 0.8059 879 221.0833
0.4380 0.6030 0.7265 651 169.7500
0.3058 0.3872 0.7899 894 223.7500
0.0343 0.1253 0.2734 1077 255.6667
0.0085 0.0574 0.1478 1101 271.4167
0.0185 0.0417 0.4437 918 234.1667
0.0220 0.0213 1.0303 930 237.6667
0.0129 0.0429 0.3014 942 239.8333
0.0122 0.0889 0.1371 675 178.8333
0.0102 0.0640 0.1588 675 179.4167
0.1969 0.4083 0.4822 612 162.5000
0.1860 0.4155 0.4476 612 163.2500
0.1886 0.4501 0.4190 612 162.7500
0.0067 0.0383 0.1744 789 187.5000
0.0100 0.0380 0.2636 792 188.9167
0.0346 0.0674 0.5137 1137 279.4167
0.0237 0.0402 0.5892 1140 281.3333

图8

16个GRF基因对外源激素响应的表达模式分析 *表示处理间存在显著差异(P < 0.05)。"

图9

16个GRF基因对环境胁迫响应的表达模式分析 *表示处理间存在显著差异(P < 0.05)。"

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