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作物学报 ›› 2025, Vol. 51 ›› Issue (6): 1514-1525.doi: 10.3724/SP.J.1006.2025.41076

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

小麦蔗糖合酶基因TaSUS2调控籽粒淀粉合成及品质的功能研究

吴美娟1,2(), 张寅辉2, 李元昊2, 刘海霞2, 黄以琳2, 李甜2, 刘红霞2, 张学勇2, 郝晨阳1,2,*(), 郭杰1,*(), 侯健2,*()   

  1. 1山西农业大学农学院/农业农村部有机旱作农业重点实验室(部省共建), 山西晋中 030801
    2中国农业科学院作物科学研究所/作物基因资源与育种全国重点实验室/农作物基因资源与基因改良国家重大科学工程, 北京 100081
  • 收稿日期:2024-11-10 接受日期:2025-03-26 出版日期:2025-06-12 网络出版日期:2025-03-31
  • 通讯作者: *郝晨阳, E-mail: haochenyang@caas.cn;郭杰, E-mail: nxgj1115326@sxau.edu.cn;侯健, E-mail: houjian@caas.cn
  • 作者简介:E-mail: 18827115116@163.com
  • 基金资助:
    本研究由国家重点研发计划项目(2023YFD1200404)

Functional dissection of sucrose synthase gene TaSUS2 regulating grain starch synthesis and quality in wheat

WU Mei-Juan1,2(), ZHANG Yin-Hui2, LI Yuan-Hao2, LIU Hai-Xia2, HUANG Yi-Lin2, LI Tian2, LIU Hong-Xia2, ZHANG Xue-Yong2, HAO Chen-Yang1,2,*(), GUO Jie1,*(), HOU Jian2,*()   

  1. 1College of Agronomy, Shanxi Agricultural University / Key Laboratory of Sustainable Dryland Agriculture (Co-construction by Ministry and Province), Ministry of Agriculture and Rural Affairs, Jinzhong 030801, Shanxi, China
    2Institute of Crop Sciences, Chinese Academy of Agricultural Sciences / State Key Laboratory of Crop Gene Resources and Breeding / National Key Facility for Crop Gene Resources and Genetic Improvement, Beijing 100081, China
  • Received:2024-11-10 Accepted:2025-03-26 Published:2025-06-12 Published online:2025-03-31
  • Contact: *E-mail: haochenyang@caas.cn;E-mail: nxgj1115326@sxau.edu.cn;E-mail: houjian@caas.cn
  • Supported by:
    the National Key Research and Development Program of China(2023YFD1200404)

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摘要：

小麦是重要的粮食作物之一, 持续提升其产量是育种的重要目标。提高粒重可以有效提高小麦单产, 而小麦籽粒的主要成分是淀粉。为全面解析淀粉合成通路关键酶基因TaSUS2对籽粒淀粉合成的作用, 本研究从小麦基因组中扩增了TaSUS2的全长cDNA序列, 并在科农199中对TaSUS2进行基因编辑, 获得2种纯合二突材料(KO-1、KO-2)和1种纯合的三突材料(KO-3)。转基因材料表型鉴定发现, 与野生型相比, TaSUS2的种子出现明显的皱缩, 粒重显著下降, 且籽粒胚乳中总淀粉含量、直链淀粉含量、绝对淀粉含量和A型淀粉的粒径也显著降低, 表明TaSUS2影响粒重和籽粒中的淀粉合成。转录组分析发现, TaSUS2-KO-3花后21 d的籽粒中淀粉合成通路上的多个主要合成酶基因上调表达。开发TaSUS2-2A-CAPS标记并鉴定了145份重测序材料, 发现TaSUS2影响淀粉含量、湿面筋含量、蛋白质含量及沉降值等品质性状, TaSUS2-2A-Hap-G是影响品质性状的优异单倍型。研究结果为阐析TaSUS2的生物学功能奠定了基础, 也为未来小麦的高产优质分子育种提供了基因资源。

关键词: 小麦, TaSUS2, 粒重, 淀粉含量, 品质

Abstract:

Wheat is one of the world’s most important cereal crops, and improving yield remains a key goal in wheat breeding. Grain weight is a major determinant of yield, and starch is the primary component of wheat grains. To investigate the function of TaSUS2, a key enzyme gene in the starch synthesis pathway, we amplified its full-length cDNA sequence from the wheat genome and performed gene editing in the cultivar Kenong 199 (KN199). This resulted in the generation of two homozygous diploid mutants (KO-1 and KO-2) and one homozygous triploid mutant (KO-3). Phenotypic analysis of the transgenic lines revealed that TaSUS2 mutant grains exhibited pronounced wrinkling and a significant reduction in grain weight compared to the wild type. Additionally, the total starch content, amylose content, absolute starch content, and the diameter of A-type starch granules in the endosperm were significantly reduced in TaSUS2 mutant grains. These findings confirm that TaSUS2 plays a crucial role in starch synthesis and grain weight determination. Transcriptome analysis indicated that multiple enzyme-encoding genes involved in starch biosynthesis were upregulated in TaSUS2-KO-3 grains at 21 days post-anthesis (DPA). Furthermore, genotyping of a natural population of 145 wheat accessions using the TaSUS2-2A-CAPS marker revealed that TaSUS2 was significantly associated with starch content, wet gluten content, protein content, and sedimentation value. Notably, the TaSUS2-2A-Hap-G haplotype was identified as a favorable allele for these quality traits. Overall, this study provides valuable insights into the biological function of TaSUS2 and offers novel genetic resources for molecular breeding aimed at improving wheat yield and quality.

Key words: wheat, TaSUS2, grain weight, starch content, quality

表1

本试验所用的引物"

引物用途
Primer use		 引物名称
Primer name		 引物序列
Primer sequence (5′-3′)
基因扩增 Gene amplification SUS2-2A-F ATGAACAAACGCGGCT
SUS2-2A-R TCATTTGCCCGAGGTC
SUS2-2B-F ATGATCTGCTTCCCAAAGCA
SUS2-2B-R TCATTTGCCCGAGGTC
CAPS标记 CAPS marker SUS2-2A-M1F GCAATAGTTCCGTGCTCCTGTG
SUS2-2A-M1R AGAAATACGCAAGGCAACCAT
SUS2-2A-M2F CACGCCAACCTTTCTTCTTCC
SUS2-2A-M2R TCGCTTAACCGTGTCTCACA

图1

TaSUS2的编辑位点 TaSUS2在科农199中的编辑位点。WT(野生型)展示了TaSUS2第3外显子上sgRNA的靶向区域, PAM序列用红色突出显示。KO-1、KO-2、KO-3表示3种突变体的基因型, 后面的“+”和“-”分别表示由CRISPR/Cas9编辑引起的碱基插入或缺失, 数字表示插入或缺失的碱基数目。序列最右侧的数字表示该基因翻译的氨基酸数目。"

图2

TaSUS2-KO籽粒的表型及品质性状 A: WT和TaSUS2-KO小麦籽粒的形态。标尺为5 mm。B~F: WT和TaSUS2-KO小麦的千粒重、粒长、粒宽、粒厚、单株产量比较。G: WT和TaSUS2-KO花后20 d胚乳中的淀粉颗粒形态和淀粉合成情况。标尺为20 μm。H: WT和TaSUS2-KO小麦籽粒的总淀粉含量。I: WT和TaSUS2-KO小麦籽粒的直链淀粉含量。J: WT和TaSUS2-KO小麦籽粒中的支链淀粉含量。K: WT和TaSUS2-KO小麦籽粒中的绝对淀粉含量。L: WT和TaSUS2-KO胚乳中A型淀粉的粒径。M: WT和TaSUS2-KO小麦籽粒中的总蛋白含量。**: P < 0.01; ***: P < 0.001; ns表示差异不显著。"

图3

TaSUS2-KO籽粒的转录组分析 A: WT与TaSUS2-KO-3花后7、14、21 d籽粒转录组差异基因数量比较。B: 花后21 d TaSUS2-KO-3主要差异表达基因GO富集。C: 花后21 d TaSUS2-KO-3淀粉合成通路主要合成酶变化特征。DPA: 花后天数。SUS: 蔗糖合酶; FRK: 果糖激酶; PGI: 磷酸葡萄糖异构酶; PGM: 磷酸葡萄糖变位酶; UGPase: UDP-葡萄糖焦磷酸化酶; AGPase: ADP-葡萄糖焦磷酸化酶; BT1: ADP-葡萄糖转运蛋白; GBSSI: 造粉体内淀粉合成相关酶如颗粒结合型淀粉酶I; SS: 淀粉合酶; SBE: 淀粉分支酶; ISA: 异淀粉酶; PUL: 极限糊精酶。"

表2

TaSUS2-KO-3花后21 d籽粒中生长素信号通路基因差异表达分析"

生长素信号通路基因ID
Auxin signalling pathway gene ID		 生长素信号通路基因注释
Gene annotation of auxin signaling pathway		 差异倍数
Difference multiple
TraesCS2D02G322600 ABC transporter B family member 19 0.39
TraesCS2A02G344300 ABC transporter B family member 19 0.32
TraesCS2B02G341800 ABC transporter B family member 19 0.24
TraesCS2D02G548900 Auxin response factor 12 0.37
TraesCS2A02G547800 Auxin response factor 12 0.46
TraesCS2B02G578500 Auxin response factor 12 0.34
TraesCS7A02G252000 Auxin response factor 21 0.45
TraesCS7B02G138900 Auxin response factor 21 0.43
TraesCS7D02G250100 Auxin response factor 21 0.46
TraesCS5D02G045700 Auxin response factor 25 0.36
TraesCS5B02G039800 Auxin response factor 25 0.37
TraesCS5A02G038300 Auxin response factor 25 0.42
TraesCS6A02G138600 Auxin response factor 6 0.38
TraesCS6D02G127600 Auxin response factor 6 0.35
TraesCS6B02G167100 Auxin response factor 6 0.24
TraesCS1A02G077800 Auxin transporter-like protein 3 0.28
TraesCS1D02G079900 Auxin transporter-like protein 3 0.33
TraesCS1B02G095900 Auxin transporter-like protein 3 0.31
TraesCS5B02G104100 Auxin-binding protein 1 0.39
TraesCS5A02G098900 Auxin-binding protein 1 0.41
TraesCS3B02G181500 Auxin-responsive protein IAA17 0.44
TraesCS7A02G331100 Auxin-responsive protein IAA21 0.47
TraesCS3A02G155200 Auxin-responsive protein IAA3 0.48
TraesCS5D02G069300 Auxin-responsive protein IAA30 0.35
TraesCS5B02G058500 Auxin-responsive protein IAA30 0.31
TraesCS5A02G058600 Auxin-responsive protein IAA30 0.40
TraesCS7A02G384600 Auxin-responsive protein SAUR32 0.47
TraesCS1A02G178700 Auxin-responsive protein SAUR32 0.43
TraesCS6A02G141200 Auxin-responsive protein SAUR71 0.15
TraesCS6B02G169500 Auxin-responsive protein SAUR71 0.43
TraesCS4B02G016500 Auxin-responsive protein SAUR71 0.31
TraesCS7D02G191600 Probable auxin efflux carrier component lc 0.33
TraesCS7A02G190600 Probable auxin efflux carrier component lc 0.37

表3

TaSUS2-KO-3花后21 d籽粒中淀粉合成相关酶基因(SSRG)差异表达分析"

淀粉合成相关酶基因
SSRG		 差异倍数
Difference multiple		 淀粉合成相关酶基因
SSRG		 差异倍数
Difference multiple		 淀粉合成相关酶基因
SSRG		 差异倍数
Difference multiple
SUS2-2A 0.09 AGPL-1B 1.95 SSIIb-6D 0.90
SUS2-2B 0.14 AGPL-1D 2.10 SSIII-1A 1.89
SUS2-2D 0.17 AGPS1-7A 1.51 SSIII-1B 2.22*
SUS1-7A 1.47 AGPS1-7B 2.98* SSIII-1D 1.76
SUS1-7B 1.25 AGPS1-7D 2.50* SBEI-7A 0.79
SUS1-7D 1.33 BT1-6A 2.35* SBEI-7B 1.06
UGP-5A 1.25 BT1-6B 2.03* SBEI-7D 0.92
UGP-5B 1.31 BT1-6D 2.73* SBEIIa-2A 2.11*
UGP-5D 1.19 GBSSI-4A 1.52 SBEIIa-2B 1.90
FRK-7A 1.55 GBSSI-7A 1.98 SBEIIa-2D 1.72
FRK-7B 1.68 GBSSI-7D 1.65 SBEIIb-2A 2.13*
FRK-7D 1.37 SSI-7A 1.60 SBEIIb-2B 2.18*
PGI-1A 0.92 SSI-7B 1.48 SBEIIb-2D 2.57*
PGI-1B 1.31 SSI-7D 1.79 ISA-7A 1.58
PGI-1D 1.28 SSIIa-7A 2.73* ISA-7B 1.66
PGM-4A 1.22 SSIIa-7B 2.02* ISA-7D 1.39
PGM-4B 1.33 SSIIa-7D 2.37* PUL-7A 1.01
PGM-4D 1.28 SSIIb-6A 0.75 PUL-7B 1.64
AGPL-1A 1.47 SSIIb-6B 0.62 PUL-7D 1.96

图4

TaSUS2-2A-CAPS标记开发与单倍型/品质性状的相关性分析 A: TaSUS2-2A-CAPS标记开发。TaSUS2-2A-MF/R的PCR产物被Asc I酶切后, TaSUS2-2A-Hap-A无法被切开, TaSUS2-2A-Hap-G产物被切为175 bp+166 bp。B~G: TaSUS2-2A不同单倍型与品质性状在145份自然群体中的相关性分析。B: 总淀粉含量; C: 直链淀粉含量; D: 容重; E: 蛋白质含量; F: 湿面筋含量; G: 沉降值。ns表示差异不显著。"

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