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作物学报 ›› 2023, Vol. 49 ›› Issue (3): 597-607.doi: 10.3724/SP.J.1006.2023.22014

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

包穗突变体sui1-5鉴定及OsPSS1互作蛋白筛选

杨晔1(), 孙琦2, 邢欣欣2, 张海涛1,*(), 赵志超2,*(), 程治军2   

  1. 1安徽农业大学农学院, 安徽合肥 230036
    2中国农业科学院作物科学研究所, 北京 100081
  • 收稿日期:2022-03-05 接受日期:2022-07-21 出版日期:2023-03-12 网络出版日期:2022-08-19
  • 通讯作者: 张海涛,赵志超
  • 作者简介:E-mail: 874498704@qq.com
  • 基金资助:
    国家自然科学基金项目(31871603)

Identification of sheathed panicle mutant sui1-5 and screening of OsPSS1 interaction protein in rice (Oryza sativa L.)

YANG Ye1(), SUN Qi2, XING Xin-Xin2, ZHANG Hai-Tao1,*(), ZHAO Zhi-Chao2,*(), CHENG Zhi-Jun2   

  1. 1College of Agronomy, Anhui Agricultural University, Hefei 230036, Anhui, China
    2Institute of Crop Sciences, Chinese Academy of Agricultural Sciences, Beijing 100081, China
  • Received:2022-03-05 Accepted:2022-07-21 Published:2023-03-12 Published online:2022-08-19
  • Contact: ZHANG Hai-Tao,ZHAO Zhi-Chao
  • Supported by:
    National Natural Science Foundation of China(31871603)

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摘要：

包穗是水稻生产上的不利性状, 表现为倒一节间伸长异常, 穗部不能正常从叶鞘伸出。目前, 已经鉴定报道了多个包穗性状突变体, 但对该性状产生的分子机制认识仍然有限。OsPSS1/SUI1是一个磷脂酰丝氨酸合酶的编码基因, 该基因突变会引起突变体sui1-4的倒一节间严重缩短。我们鉴定到了一个新的OsPSS1/SUI1等位突变体sui1-5, 该突变体倒一节间缩短程度明显减轻。测序发现突变体中基因SUI1的第7个外显子存在1个单碱基G→T替换, 导致甘氨酸突变为缬氨酸。为了研究OsPSS1的功能, 我们通过酵母筛库发现了一个与OsPSS1互作的蛋白GH9A3, 并通过酵母双杂交、荧光素酶互补(LCI)和双分子荧光互补(BIFC)实验加以验证。GH9A3是GH9基因家族成员, 编码内切-1,4-β葡聚糖酶, 推测是细胞壁成分纤维素合成的一个关键酶。在苗期, SUI1和互作蛋白编码基因GH9A3的表达量并不同步, GH9A3在sui1-5的表达量上调。但是在拔节抽穗期, SUI1和GH9A3在sui1-5中的表达量都相应降低, 其他纤维素合酶CESAs家族基因的表达量在sui1-5中也明显降低。这些结果为进一步研究OsPSS1的功能奠定了一定的理论基础。

关键词: 包穗, SUI1, GH9A3, 精细定位, 纤维素合酶

Abstract:

The sheathed panicle phenomenon is a disadvantageous trait in agricultural production, which is mainly due to the abnormal elongation of the uppermost internode, resulting in the panicle closed by flag leaf sheath. Although some mutants of sheathed panicle have been identified and reported in rice, the underling molecular mechanism of uppermost internode shortening is still poorly understood. OsPSS1/SUI1 is a phosphatidylserine synthase gene and the mutation of OsPSS1 leads to a severe shortening of the uppermost internode of mutant sui1-4. We discovered and identified a new allelic mutant sui1-5 of SUI1 and the degree of the uppermost internode shortening in sui1-5 was significantly reduced. Sequence analysis revealed a single nucleotide substitution of G to T at the seventh exon of SUI1 in sui1-5, resulting in an amino acid change from glycine to valine. To study the function of OsPSS1, we detected a protein GH9A3 interacting with SUI1 through a yeast screening library, which was verified by yeast two-hybrid, luciferase complementarity (LCI), and bimolecular fluorescence complementarity (BiFC). GH9A3 was a member of GH9 gene family, encoding an endo-β-1,4-glucanase which was speculated to be a key enzyme in cell wall cellulose synthesis. At seedling stage, the relative expression level of SUI1 was not synchronized with that of interaction protein-coding gene GH9A3, and the relative expression level of GH9A3 was up-regulated in sui1-5. But at heading and jointing stage, the relative expression level of SUI1 and GH9A3 decreased synchronously, and the relative expression level of other cellulose synthase CESAs family genes decreased obviously in sui1-5. These results lay a foundation for further study of the function of OsPSS1.

Key words: sheathed panicle, SUI1, GH9A3, fine mapping, cellulose synthesis

表1

用于基因精细定位的Indel和SNP引物"

多态性分子标记
Polymorphic marker		 正向引物
Forward sequence (5'-3')		 反应引物
Reverse sequence (5'-3')
Inc1-1 AGGGGAAGAAAAACCTGACC CCGCGTGCAGATAAAGTACA
Y1-1 CAAGCAGTGATCATACAGCCTTCC GCCATGGCTGAGAACAGAGAGC
Y1-2 CTCCTGTTACAACTGCATGGA TCGATCCATTAGCTAGCCGG
Y1-3 CATGAGTACAACATCGCCTACA CGATGATTCAGTACATGCATGC
Y1-4 GCTGCATCGGTATACAAGACG TCAGTTTGTCAGGGAGGGAG
Y1-5 CTCCGTTCTCCACCGTGT AACAGCCAACTAGGTTCACA
Y1-6 GCGGCCACCATGATCTTGTACC GTACTTGTTGTAGCGCCCGTTCC
Y1-7 CTAGCAGCTGTCTGCGACACACG CCGAGGTGTTATGCCAATCTCTATGG

表2

用于qRT-PCR引物"

基因名称
Gene name		 正向引物
Forward sequence (5'-3')		 反应引物
Reverse sequence (5'-3')
OsCesA1 TCTTCTCGCCACGAACAACA CTTGGAGGGGTCCACAATCC
OsCesA3 TGCCAGCCATCTGTTTGCTA GCCAACACCACTCCATCTCA
OsCesA4 AAGGGATCAGCGCCAATCAA GGAGCCATTTCAGACGACCA
OsCesA7 TGGAAGTCGGTGTACTGCAC GCGGCTCATGAAGATCTCGA
OsCesA8 ATGACCAACGGGACAAGCAT TAGAGAGAGGCTGGCGAGTT
OsCesA9 GCGCCGATCAACCTATCTGA CTTGTAGCCGTAGAGGAGCG
OsGH9A3 TGAGTGCCACTCTTGCTCAG CCGAACTTGTACAGGGCCTT
SUI1 GCAGACCCGTGATGATGCTA TATATGCGGCAGTCGGTTCC
Ubiquitin ACCACTTCGACCGCCACTACT ACGCCTAAGCCTGCTGGTT

图1

野生型和突变体表型对比 A: 野生型Kitaake (左)与突变体sui1-4 (右)抽穗期表型对比; B: 野生型Kitaake (左)与突变体sui1-5 (右)抽穗期表型对比; C: 野生型(左)和突变体(右)穗部表型以及各节间长度对比; D: 野生型与突变体各节间长度对株高的贡献率; E, F: 野生型(E)和突变体(F)倒二节间纵切面细胞学比较; G, H: 野生型和突变体纵切薄壁细胞长度和宽度; I, J: 野生型(I)和突变体(J)倒二节间横切面细胞学比较; K: 野生型和突变体倒二节间横截面半径细胞数目。标尺: (A) 10 cm, (B) 10 cm, (C) 10 cm, (E, F) 100 μm, (I, J) 100 μm。*表示在P < 0.05水平差异显著。"

表3

野生型和突变体农艺性状比较"

农艺性状
Agronomic trait		 野生型
Kitaake		 突变体
sui1-5
株高Plant height (cm) 78.90±4.80 68.80±5.40**
分蘖数Tiller number 23.40±2.37 31.50±3.30**
剑叶宽Flag leaf length (cm) 1.21±0.09 1.08±0.08**
一次枝梗Primary branch number 6.80±0.92 8.30±1.50*
二次枝梗Secondary branch number 7.50±1.96 1.70±1.25**
穗长 panicle length (cm) 14.09±0.96 9.27±1.01**
倒一节间长First internode length (cm) 26.64±3.47 0.21±0.12**
倒二节间长Second internode length (cm) 17.51±0.98 14.18±0.88**
倒三节间长Third internode length (cm) 9.35±1.19 8.63±1.58
倒四节间长Fourth internode length (cm) 2.53±0.37 1.79±0.51**
每穗粒数Grain number per panicle 67.10±8.78 29.80±4.64**
结实率Seed setting rate (%) 96.00±0.03 62.00±0.06**
千粒重1000-grain weight (g) 26.13±0.04 26.89±0.06**
粒长Seed length (mm) 6.79±0.15 6.87±0.19
粒宽Seed weight (mm) 3.33±0.04 3.39±0.13
粒厚Seed thickness (mm) 2.20±0.05 2.34±0.32

图2

SUI1基因的图位克隆 A: SUI1初步定位在1号染色体标记Y1-1与Y1-7之间; B: SUI1精细定位在标记Y1-3和Y1-4之间38.05 kb范围内; 横线上方为基因定位所用分子标记, 横线下方为交换单株数。C: SUI1精细定位区间内包含3个候选基因; D: SUI1基因包含12个外显子和11个内含子, 白方块表示编码区, 黑方块表示非编码区, 线条表示内含子下图为Kitaake和sui1-5在第7个外显子上的序列对比, 箭头指向突变的碱基。"

图3

SUI1与GH9A3蛋白互作验证 A: 酵母双杂交验证SUI1和GH9A3的互作。AD: 转录激活域; BD: DNA结合结构域; SD: 营养缺陷型培养基。B: 荧光素酶互补成像(LCI)验证SUI1与GH9A3在本氏烟草中的互作。C: BiFC在本氏烟草中验证SUI1与GH9A3的互作, mCherry: SCAMP1。至少重复3个独立烟草叶片。"

图4

水稻GH9A3蛋白进化树分析 水稻OsGH9A3蛋白序列作为Qurry序列, 普通小麦、玉米和拟南芥作为候选物种, 以BLAST得到的同源蛋白在MEGA7.0中采用邻接法构建进化树。引导程序值以节点上的百分比显示。分支长度与用于推断系统发育树的进化距离。结果表明OsGH9A3与玉米gpm617蛋白同源性最接近, 水稻和拟南芥GH9A家族蛋白之间同源性也比较高。"

图5

GH9A3烟草表皮亚细胞定位 上图为35S-GFP空载体对照; 下图为连接GH9A3片段的35S-GFP载体, mCherry: AtSYP122, GH9A3定位在烟草表皮细胞的质膜上, 伴随点状结构。"

图6

SUI1与GH9A3表达量在抽穗期的突变体中显著降低 A: 幼苗期野生型和突变体sui1-5表型, 左: 野生型, 右: 突变体sui1-5; B: 基因SUI1和GH9A3在苗期(左)和抽穗期(右)表达量的变化情况。在突变体中2个基因在抽穗期的表达量均大幅下降。**表示在P < 0.01水平差异显著。"

图7

CESAs纤维素合酶家族基因表达量分析 CESAs的表达量在突变体sui1-5抽穗期的茎中显著下调。OsCesA1、OsCESA3和OsCESA8参与初级细胞壁合成; OsCesA4、OsCESA7和OsCESA9参与次级细胞壁合成。**表示在P < 0.01水平差异显著。"

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