水稻粉质胚乳突变体ws的表型分析及基因克隆

doi:10.16819/j.1001-7216.2016.6048

中国水稻科学 ›› 2016, Vol. 30 ›› Issue (5): 447-457.DOI: 10.16819/j.1001-7216.2016.6048

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水稻粉质胚乳突变体ws的表型分析及基因克隆

潘鹏屹, 朱建平, 王云龙, 郝媛媛, 蔡跃, 张文伟, 江玲, 王益华, 万建民^*()

南京农业大学作物遗传与种质创新国家重点实验室/农业部长江中下游粳稻生物学与遗传育种重点实验室/长江流域杂交水稻协同创新中心/江苏省现代作物生产中心,南京210095

收稿日期:2016-03-19 修回日期:2016-04-28 出版日期:2016-09-10 发布日期:2016-09-10
通讯作者: 万建民
基金资助:
国家转基因生物新品种培育重大专项(2014ZX08001006);国家自然科学基金资助项目(31371598);江苏省科技支撑计划资助项目(BE2014394);农业部长江中下游粳稻生物学与遗传育种重点实验室、长江流域杂交水稻协同创新中心、江苏省现代作物生产中心资助项目

Phenotyping and Gene Cloning of a Floury Endosperm Mutant ws in Rice

Peng-yi PAN, Jian-ping ZHU, Yun-long WANG, Yuan-yuan HAO, Yue CAI, Wen-wei ZHANG, Ling JIANG, Yi-hua WANG, Jian-min WAN^*()

State Key Laboratory of Crop Genetics and Germplasm Enhancement, Nanjing Agricultural University/Key Laboratory of Biology, Genetics and Breeding of japonica Rice in Mid-lower Yangtze River, Ministry of Agriculture/The Yangtze River Valley Hybrid Rice Collaboration Innovation Center/Jiangsu Collaborative Innovation Center for Modern Crop Production, Nanjing 210095, China

Received:2016-03-19 Revised:2016-04-28 Online:2016-09-10 Published:2016-09-10
Contact: Jian-min WAN

摘要/Abstract

摘要：

从甲基亚硝基脲(1-Methyl-1-Nitrosourea, MNU)处理的粳稻品种滇粳优1号突变体库中,筛选到一个稳定遗传的胚乳粉质突变体ws,其籽粒的千粒重、籽粒大小、总淀粉含量、直链淀粉含量等指标均降低,淀粉在尿素溶液中的膨胀能力减弱。对成熟及发育中的胚乳淀粉结构进行观察,发现ws突变体的胚乳中产生大量小而不规则排布的单淀粉颗粒。利用F₂群体中分离出的92个隐性极端个体将突变基因连锁在第8染色体近着丝粒位置,随后共用2025个极端个体将目标基因定位于95 kb的区间。测序发现ws突变体中编码腺苷二磷酸葡萄糖焦磷酸化酶(Adenosine diphosphate glucose pyrophosphorylase, AGPase)小亚基S2的基因发生点突变,导致编码氨基酸的替换。基因表达分析发现,突变体胚乳中编码AGPase各亚基的相关基因表达量没有发生显著改变,而Western杂交分析显示突变体中AGPS2b的蛋白含量下降。同时,ws突变体的胚乳中AGPase活性下降为野生型的一半。研究结果表明,OsAGPS2的突变导致水稻胚乳中AGPase活性降低,从而影响了淀粉合成。

关键词: 胚乳粉质突变体, AGPase, 淀粉, 基因表达, 理化性质, 水稻

Abstract:

In this study, we obtained a stably inherited floury endosperm mutant ws from japonica variety cv. Dianjingyou 1 (DJY), with its 1000-grain weight, grain size, total starch content, and amylose content in mature seeds all decreased compared with wild-type, as well as the swelling power of starch. By observing the structure of mature and developing endosperm, we found that the starch granules in ws mutant were smaller and loosely packed. The WS locus was first mapped to a region near the centromere in chromosome 8 with 92 recessive individuals, and then was narrowed down to a 95 kb region by using 2025 recessive individuals. Sequence analysis revealed that the coding region of the small subunit of adenosine diphosphate glucose pyrophosphorylase (AGPS2) had a single nucleotide substitution, resulting in a change in the amino acid sequence. RT-PCR analysis showed no significant difference in the expression levels of genes encoding the AGPase subunits in the mutant endosperm, while immunoblot analysis revealed a reduced protein content of the AGPS2b. Meanwhile, the enzyme activity of AGPase in the mutant was decreased to half of the wild-type. These results showed that the mutation of OsAGPS2 caused the decreased activity of AGPase, thus affecting the starch biosynthesis in rice endosperm.

Key words: floury endosperm mutant, AGPase, starch, gene expression, physicochemical property, rice (Oryza sativa L.)

中图分类号:

潘鹏屹, 朱建平, 王云龙, 郝媛媛, 蔡跃, 张文伟, 江玲, 王益华, 万建民. 水稻粉质胚乳突变体ws的表型分析及基因克隆[J]. 中国水稻科学, 2016, 30(5): 447-457.

Peng-yi PAN, Jian-ping ZHU, Yun-long WANG, Yuan-yuan HAO, Yue CAI, Wen-wei ZHANG, Ling JIANG, Yi-hua WANG, Jian-min WAN. Phenotyping and Gene Cloning of a Floury Endosperm Mutant ws in Rice[J]. Chinese Journal OF Rice Science, 2016, 30(5): 447-457.

图/表 9

表1 WS基因定位引物的序列

Table 1 Markers for fine mapping of WS.

标记 Marker	正向引物Forward (5'→3')	反向引物Reverse (5'→3')
HY8-19	TTTGTTGCTTTTCTGATTC	ATGATAAAGCGATAAACCA
WQ8-28	GAGACGGACGGGTGTTGA	CAATGACATCCCAGCGTA
WZ8-17	TAAATCATGGTGGTGGGC	ACCGTCGTCTAGCAAGGAG
WZ8-3	ATTAAGATGATATGGGAAGT	ACATTGACCTGGTAGAAAC
HY8-24	ATTAAGATGATATGGGAAGT	ACATTGACCTGGTAGAAAC

表2 实时RT-PCR引物序列

Table 2 Primers used in real-time RT-PCR.

基因 Gene	正向引物 Forward (5'→3')	反向引物 Reverse (5'→3')
AGPL1	CATCAAGGACGGGAAGGTCA	ACTTCACTCGGGGCAGCTTA
AGPL2	CTGAGGAAGAGGTGCTTTGG	TCTTTCGGGAGGATTGTGTC
AGPS1	AGAATGCTCGTATTGGAGAAAATG	GGCAGCATGGAATAAACCAC
AGPS2a	ACTCCAAGAGCTCGCAGACC	GCCTGTAGTTGGCACCCAGA
AGPS2b	AACAATCGAAGCGCGAGAAA	GCCTGTAGTTGGCACCCAGA
UGPase1	CCATCACCGCCAAGTCA	GACCGTTGATGTCCTTGTTCT
Actin	CCCTCCTGAAAGGAAGTACAGTGT	GTCCGAAGAATTAGAAGCATTTCC

图1 野生型与ws突变体的表型比较 A-野生型与ws突变体种子表型比较,标尺为3 mm; B-野生型与ws突变体种子横切面,标尺为1 mm; C-野生型与ws突变体植株,标尺为20 cm。DJY-滇粳优1号(野生型)。

Fig. 1. Phenotype comparison of the wild-type and ws mutant. A, Comparison of wild-type and ws mutant seeds, bar = 3 mm; B, Seed cross-sections of wild-type and ws mutant, bar = 1 mm; C, Comparison of wild-type and ws mutant plants, bar = 20 cm. DJY, Dianjingyou 1(wild type).

图2 野生型与ws突变体的千粒重、籽粒及主要农艺性状比较千粒重n = 3,其余n = 10,取平均值± SD; 采用t测验,**表示P < 0.01。DJY-滇粳优1号(野生型)。

Fig. 2. Comparison of 1000-grain weight, seed and major agronomic traits of wild-type and ws mutant. Values are mean ± SD (n = 10, except for 1000-grain weight, n = 3); t-test, ** P < 0.01. DJY, Dianjingyou 1(wild type).

图3 野生型与ws突变体成熟籽粒的理化性质分析 A-野生型与ws突变体籽粒的化学成分分析(Mean± SD),n = 3,**,P < 0.01(t测验); B-野生型与ws突变体的尿素膨胀; C-4 mol/L的尿素浓度下出现显著差异; D-不同尿素浓度下野生型和突变体的米粉的膨胀体积比较(n = 3)。DJY-滇粳优1号(野生型)。

Fig. 3. Physicochemical characteristics of wild-type and ws mature seeds. A, Comparison of chemical composition of wild-type and ws seeds, values are mean ± SD, n = 3; t-test, ** P < 0.01; B, Gelatinization properties of wild type and ws mutant seeds; C, Significant difference was observed at the urea concentration of 4 mol/L urea; D, The swollen volume of wild-type and ws starch in urea solutions of various concentrations (n = 3). DJY, Dianjingyou 1(wild type).

图4 野生型与ws突变体成熟籽粒的扫描电镜观察 A,B-野生型胚乳; C,D-ws突变体胚乳。A,C为籽粒横切面,标尺为1 mm; B,D是局部放大后的胚乳淀粉结构(红色方框内),标尺为10 μm。

Fig. 4. Scanning electron microscopy observation of mature seeds of wild type and ws mutant. A, B, Endosperm of wild-type; C, D, Endosperm of ws mutant; A, C, Endosperm cross-section, Bars = 1 mm; B, D, Partial enlarged drawing picture, Bars = 10 μm.

图5 野生型与ws突变体胚乳的半薄切片观察 A,B-开花后9 d野生型胚乳细胞外围、里层; C-开花后12 d野生型胚乳细胞里层; D,E-开花后9 d的ws突变体胚乳细胞外围、里层; F-开花后12 d的ws突变体胚乳细胞里层。A,D标尺是100 μm,B,C,E,F标尺是50 μm。

Fig. 5. Semi-thin sections of wild type and ws mutant seeds. A, B, Peripheral part (A) and central part (B) of wild-type endosperm cells at 9 DAF (days after flowering); C, Central part of wild-type endosperm cells at 12 DAF; D, E, Peripheral part (D) and central part (E) of ws endosperm cells at 9 DAF; F, Central part of ws endosperm cells at 12 DAF. Bars = 100 μm in A and D, 50 μm in B, C, E, F.

图6 WS基因的精细定位 A-WS基因与标记HY8-19和HY8-24连锁; B-利用2025个极端个体将WS基因定位在95 kb区间内; C-WS候选基因分析; D-突变体AGPS2中发生单碱基替换(红色标出),AGPS2由10个外显子(黑色方框表示)和9个内含子组成,AGPS2a与AGPS2b的第一个外显子分别为1a与1b。

Fig. 6. Fine-mapping of WS gene. A, WS was linked with markers HY8-19 and HY8-24; B, WS was located in a 95 kb region based on 2025 individuals; C, Candidate genes for WS; D, ws displayed a single nucleotide substitution in AGPS2 (in red),AGPS2 is composed of 10 exons (filled box) and 9 introns, the alternative use of exon 1a and 1b generates AGPS2a and AGPS2b transcripts, respectively.

图7 野生型及ws突变体中的相关基因表达及酶活分析 A-野生型不同组织中AGPS2b的表达分析; B-野生型不同发育时期胚乳中AGPS2b的表达分析; C-野生型和ws突变体发芽7 d后幼苗叶片中AGPS2a表达分析; D-野生型与ws突变体发育中胚乳的AGPase相关基因表达分析; E-野生型与ws突变体发育中胚乳的相关蛋白表达分析; F-野生型和ws突变体发育胚乳中AGPase和UGPase活性测定。A,B,C,D,F中,n = 3,取平均值± SD。

Fig. 7. Expression of related genes and enzyme activity in wild type and ws mutant. A, Real-time RT-PCR analysis of the expression of AGPS2b in different organs in wild type; B, Real-time RT-PCR analysis of the expression of AGPS2b at different developing stages of endosperm in wild type; C, Real-time RT-PCR analysis of the expression of AGPS2a in leaves of the wild type and ws mutant seedlings at 7 days after germination; D, Real-time RT-PCR analysis of the expression of genes encoding AGPase in wild type and ws mutant endosperm; E, Immunoblot analysis of starch biosynthesis related proteins in wild type and ws mutant endosperm; F, AGPase and UGPase activities of wild-type and ws developing endosperm. For A, B, C, D and F, error bars show SD (n = 3).

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水稻粉质胚乳突变体ws的表型分析及基因克隆

Phenotyping and Gene Cloning of a Floury Endosperm Mutant ws in Rice

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