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作物学报 ›› 2022, Vol. 48 ›› Issue (1): 40-47.doi: 10.3724/SP.J.1006.2022.01103

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

小麦谷氨酰胺合成酶基因可变剪接分析

韦一昊1(), 于美琴2, 张晓娇1, 王露露1, 张志勇1, 马新明1, 李会强2, 王小纯1,2,*()   

  1. 1河南省粮食作物协同创新中心 / 河南农业大学, 河南郑州 450002
    2河南农业大学生命科学学院, 河南郑州 450002
  • 收稿日期:2020-12-29 接受日期:2021-04-14 出版日期:2022-01-12 网络出版日期:2021-06-28
  • 通讯作者: 王小纯
  • 作者简介:E-mail: yc_yihao@163.com
  • 基金资助:
    国家自然科学基金项目(32071956);河南省现代农业产业技术体系建设专项资助(S2010-01-G04)

Alternative splicing analysis of wheat glutamine synthase genes

WEI Yi-Hao1(), YU Mei-Qin2, ZHANG Xiao-Jiao1, WANG Lu-Lu1, ZHANG Zhi-Yong1, MA Xin-Ming1, LI Hui-Qing2, WANG Xiao-Chun1,2,*()   

  1. 1Collaborative Innovation Center of Henan Grain Crops, Henan Agricultural University, Zhengzhou 450002, Henan, China
    2Department of Biochemistry, College of Life Science, Henan Agricultural University, Zhengzhou 450002, Henan, China
  • Received:2020-12-29 Accepted:2021-04-14 Published:2022-01-12 Published online:2021-06-28
  • Contact: WANG Xiao-Chun
  • Supported by:
    National Natural Science Foundation of China(32071956);Agriculture Research System of Henan Province(S2010-01-G04)

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摘要：

谷氨酰胺合成酶(glutamine synthetase, GS)是植物氮素同化的关键酶, 小麦(Triticum aestivum L.) GS由12个核基因编码, 即TaGS1;1-6A/6B/6D、TaGS1;2-4A/4B/4D、TaGS1;3-4A/4B/4D和TaGS2-2A/2B/2D。利用单分子测序技术获得了TaGS基因的全长转录本, 发现TaGS1;1-6A有1种可变剪接转录本、TaGS1;1-6B有2种可变剪接形式; 与正常转录本编码的TaGS相比, TaGS1;1-6A-1的Gln_synt_N结构域部分缺失, TaGS1;1-6B-3缺少Gln_synth_ gly_rich_site结构域; TaGS1;1-6B-4是新鉴定的转录本, 编码缺少Gln_synth_gly_rich_site和Gln_synth_cat_dom结构域的GS蛋白。进一步研究了氮肥对不同氮效率小麦品种可变剪接转录本表达的影响, 发现氮高效品种YM49叶片TaGS1;1-6A-1表达水平显著高于氮低效品种XN509, 且随着氮肥增加而升高; YM49根系TaGS1;1-6A-1表达随供氮量增加而升高, XN509却呈现相反趋势。TaGS1;1-6B-3在YM49和XN509中有相似的表达趋势, 高浓度NO3-促进TaGS1;1-6B-3在叶片中表达, 却抑制其在根中的表达; 高浓度NH4+抑制TaGS1;1-6B-3在叶片中表达, 但在根中表达影响较小。TaGS1;1-6B-4主要在低氮条件下表达, NH4+浓度不影响其在YM49根中的表达。了解TaGS基因可变剪接有助于阐明TaGS同工酶在小麦氮代谢中的功能。

关键词: 小麦, 谷氨酰胺合成酶, 转录, 可变剪接

Abstract:

As a key enzyme for nitrogen assimilation in plant, glutamine synthetase (TaGS) is encoded by 12 genes in common wheat (Triticum aestivum L.), namely TaGS1;1-6A/6B/6D, TaGS1;2-4A/4B/4D, TaGS1;3-4A/4B/4D, and TaGS2-2A/2B/2D. Transcripts of TaGS genes were obtained using the single molecular sequencing technology, and the results showed that TaGS1;1-6A had one alternative splicing (AS) version of transcripts and TaGS1;1-6B had two AS versions. Compared with the TaGS encoded by normal transcript, TaGS1;1-6A-1 lacked part of the Gln_synt_N domain, and TaGS1;1-6B-3 lacked the Gln_synth_gly_rich_ site domain; TaGS1;1-6B-4 was a new AS event, and encoded a truncated GS without Gln_synth_gly_rich_site and Gln_synth_ cat_dom domain. The relative expression level of TaGS alternative splicing transcripts of wheat cultivars with different nitrogen use efficiencies showed that the relative expression level of TaGS1;1-6A-1 in leaves of YM49 (N-efficient) was significantly higher than that of XN509 (N-inefficient), and increased with the increase of nitrogen supply. In roots, the relative expression of TaGS1;1-6A-1 increased with the increase of nitrogen supply in YM49, but decreased in XN509. TaGS1;1-6B-3 showed a similar expression trend in YM49 and XN509, and high concentration of NO3- promoted the expression of TaGS1;1-6B-3 in leaves while inhibited in roots. High concentration of NH4+ inhibited the expression of TaGS1;1-6B-3 in leaves, but had little effect in roots. TaGS1;1-6B-4 was mainly expressed under low nitrogen condition, and its expression in roots of YM49 was not affected by NH4+ supply. Understanding the AS transcript versions of TaGS genes is helpful to illustrate the functions of TaGS isozymes in nitrogen metabolism of wheat.

Key words: glutamine synthetase, transcription, alternative splicing, wheat, glutamine synthetase, transcription, alternative splicing

表1

不同氮处理营养液组成"

氮含量
Nitrogen content		 KH2PO4 MgSO4 KCl CaCl2 Ca(NO3-)2 NH4Cl
N0 0.2 1 1.5 2.5 0 0
0.2 mmol L-1 NO3- 0.2 1 1.5 2.4 0.1 0
2 mmol L-1 NO3- 0.2 1 1.5 1.5 1 0
5 mmol L-1 NO3- 0.2 1 1.5 0 2.5 0
10 mmol L-1 NO3- 0.2 1 1.5 0 5 0
20 mmol L-1 NO3- 0.2 1 1.5 0 10 0
0.2 mmol L-1 NH4+ 0.2 1 1.5 2.5 0 0.2
2 mmol L-1 NH4+ 0.2 1 1.5 2.5 0 2
5 mmol L-1 NH4+ 0.2 1 1.5 2.5 0 5
10 mmol L-1 NH4+ 0.2 1 1.5 2.5 0 10
20 mmol L-1 NH4+ 0.2 1 1.5 2.5 0 20

表2

RT-PCR引物"

转录本名称
Transcript name		 引物名称
Primer name		 引物序列
Primer sequence
(5′-3′)		 退火温度
Annealing temperature (℃)		 PCR产物长度
PCR fragment
(bp)
TaGS1;1-6A-1 TaGS1;1-6A-1-F GGCCAATACCCATATTTTCACTGTG 60 1199
TaGS1;1-6A-1-R CAGTGGCCACCCACCAAATC
TaGS1;1-6B-3 TaGS1;1-6B-3-F GCCGGAGTTGTCATCGTGG 65 460
TaGS1;1-6B-3-R CAGACACCCTGATGACGACG
TaGS1;1-6B-4 TaGS1;1-6B-4-F CCTCAGTCAGCCGGCCAT 59 708
TaGS1;1-6B-4-R GATGATTACACGACGAGAAGGTAGC
TaTEF1 TaTEF1-F GGTGACAACATGATTGAGAGGTCC 60 678
TaTEF1-R AACAGCCACAGTTTGCCTCATG

图1

TaGS基因在染色体上的位置分布 箭头所示位置是染色体着丝粒。"

图1

TaGS基因在染色体上的位置分布 箭头所示位置是染色体着丝粒。"

表3

TaGS转录本在IWGSC RefSeq v2.0和第3代测序中的差异"

TaGS同工酶 TaGS isoforms 基因编号
Gene ID		 IWGSC RefSeq2.0 第3代测序分析
Third-generation sequencing analysis
转录本编号
Transcript ID		 蛋白质
Protein (aa)		 分子量
MW (kD)		 转录本编号
Transcript ID		 蛋白质
Protein (aa)		 分子量
MW
(kD)		 CPM*
TaGS1;1 TraesCS6A02G298100 TraesCS6A02G298100.1 349 38.4 TraesCS6A02G298100.1 359 39.7 6.4
TraesCS6A02G298100.2 356 39.2 TraesCS6A02G298100.2 356 39.2 103.5
TraesCS6B02G327500 TraesCS6B02G327500.1 356 39.2 TraesCS6B02G327500.1 356 39.2 154.9
TraesCS6B02G327500.2 325 35.8 0
TraesCS6B02G327500.3 322 35.6 TraesCS6B02G327500.3 322 35.6 7.6
ONT.45646.3 223 24.5 3.8
TraesCS6D02G383600LC TraesCS6D02G383600LC.1 356 39.2 TraesCS6D02G383600LC.1 356 39.2 16
TraesCS6D02G383600LC.2 325 35.8 0
TaGS1;2 TraesCS4A02G063800 TraesCS4A02G063800.1 354 38.7 TraesCS4A02G063800.1 354 38.7 114.6
TraesCS4B02G240900 TraesCS4B02G240900.1 354 38.7 TraesCS4B02G240900.1 354 38.7 134.2
TraesCS4D02G240700 TraesCS4D02G240700.1 354 38.7 TraesCS4D02G240700.1 354 38.7 170
TaGS1;3 TraesCS4A02G266900 TraesCS4A02G266900.1 362 39.6 TraesCS4A02G266900.1 362 39.6 8.8
TraesCS4B02G047400 TraesCS4B02G047400.1 362 39.5 TraesCS4B02G047400.1 362 39.5 3.3
TraesCS4D02G047400 TraesCS4D02G047400.1 362 39.5 TraesCS4D02G047400.1 362 39.5 13
TaGS2 TraesCS2A02G500400 TraesCS2A02G500400.1 427 46.7 TraesCS2A02G500400.1 427 46.7 9.7
TraesCS2A02G500400.2 423 46.09 0
TraesCS2B02G528300 TraesCS2B02G528300.1 423 46.08 0
TraesCS2B02G528300.2 369 40.63 TraesCS2B02G528300.2 427 46.7 53
TraesCS2D02G500600 TraesCS2D02G500600.1 427 46.7 TraesCS2D02G500600.1 427 46.7 27.3

表3

TaGS转录本在IWGSC RefSeq v2.0和第3代测序中的差异"

TaGS同工酶 TaGS isoforms 基因编号
Gene ID		 IWGSC RefSeq2.0 第3代测序分析
Third-generation sequencing analysis
转录本编号
Transcript ID		 蛋白质
Protein (aa)		 分子量
MW (kD)		 转录本编号
Transcript ID		 蛋白质
Protein (aa)		 分子量
MW
(kD)		 CPM*
TaGS1;1 TraesCS6A02G298100 TraesCS6A02G298100.1 349 38.4 TraesCS6A02G298100.1 359 39.7 6.4
TraesCS6A02G298100.2 356 39.2 TraesCS6A02G298100.2 356 39.2 103.5
TraesCS6B02G327500 TraesCS6B02G327500.1 356 39.2 TraesCS6B02G327500.1 356 39.2 154.9
TraesCS6B02G327500.2 325 35.8 0
TraesCS6B02G327500.3 322 35.6 TraesCS6B02G327500.3 322 35.6 7.6
ONT.45646.3 223 24.5 3.8
TraesCS6D02G383600LC TraesCS6D02G383600LC.1 356 39.2 TraesCS6D02G383600LC.1 356 39.2 16
TraesCS6D02G383600LC.2 325 35.8 0
TaGS1;2 TraesCS4A02G063800 TraesCS4A02G063800.1 354 38.7 TraesCS4A02G063800.1 354 38.7 114.6
TraesCS4B02G240900 TraesCS4B02G240900.1 354 38.7 TraesCS4B02G240900.1 354 38.7 134.2
TraesCS4D02G240700 TraesCS4D02G240700.1 354 38.7 TraesCS4D02G240700.1 354 38.7 170
TaGS1;3 TraesCS4A02G266900 TraesCS4A02G266900.1 362 39.6 TraesCS4A02G266900.1 362 39.6 8.8
TraesCS4B02G047400 TraesCS4B02G047400.1 362 39.5 TraesCS4B02G047400.1 362 39.5 3.3
TraesCS4D02G047400 TraesCS4D02G047400.1 362 39.5 TraesCS4D02G047400.1 362 39.5 13
TaGS2 TraesCS2A02G500400 TraesCS2A02G500400.1 427 46.7 TraesCS2A02G500400.1 427 46.7 9.7
TraesCS2A02G500400.2 423 46.09 0
TraesCS2B02G528300 TraesCS2B02G528300.1 423 46.08 0
TraesCS2B02G528300.2 369 40.63 TraesCS2B02G528300.2 427 46.7 53
TraesCS2D02G500600 TraesCS2D02G500600.1 427 46.7 TraesCS2D02G500600.1 427 46.7 27.3

图2

TaGS可变剪接的基因结构比较"

图2

TaGS可变剪接的基因结构比较"

图3

转录本TaGS1;1-6A-1和TaGS1;1-6B-4 PCR扩增 M: DNA分子量标记。箭头所指为目的条带。"

图3

转录本TaGS1;1-6A-1和TaGS1;1-6B-4 PCR扩增 M: DNA分子量标记。箭头所指为目的条带。"

图5

不同氮处理下TaGS1;1可变剪接转录本的表达"

图5

不同氮处理下TaGS1;1可变剪接转录本的表达"

图4

TaGS转录本对应的蛋白质结构域"

图4

TaGS转录本对应的蛋白质结构域"

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