中国水稻科学 ›› 2018, Vol. 32 ›› Issue (5): 415-426.DOI: 10.16819/j.1001-7216.2018.8003
• • 下一篇
李景芳1, 田云录1, 刘喜1, 刘世家1, 陈亮明1, 江玲1, 张文伟1, 徐大勇2, 王益华1,*(), 万建民1
收稿日期:
2018-01-15
修回日期:
2018-03-17
出版日期:
2018-09-10
发布日期:
2018-09-10
通讯作者:
王益华
基金资助:
Jingfang LI1, Yunlu TIAN1, Xi LIU1, Shijia LIU1, Liangming CHEN1, Ling JIANG1, Wenwei ZHANG1, Dayong XU2, Yihua WANG1,*(), Jianmin WAN1
Received:
2018-01-15
Revised:
2018-03-17
Online:
2018-09-10
Published:
2018-09-10
Contact:
Yihua WANG
摘要:
【目的】对水稻粉质皱缩突变体fse2进行表型分析及基因克隆,为阐明水稻淀粉合成机制以及胚的发育奠定基础。【方法】fse2来自粳稻品种滇粳优1号的MNU(N-甲基-N-亚硝基脲)诱变突变体库。本研究考查了突变体fse2籽粒的理化性状,利用扫描电镜和半薄切片观察了淀粉颗粒的结构;构建了fse2与N22的F2群体,通过图位克隆及转基因互补验证确定目标基因;通过qRT-PCR以及GUS活性染色对FSE2进行组织表达分析;免疫印迹分析了突变体中淀粉合成相关基因以及线粒体基因的蛋白变化。【结果】fse2籽粒粉质皱缩,千粒重显著下降;胚乳中淀粉颗粒变小变圆,排列松散,不能形成正常的复合淀粉颗粒;突变体中总淀粉、直链淀粉含量均显著下降,脂肪含量显著上升,突变体淀粉的糊化特性发生明显改变。FSE2编码一个线粒体和质体双定位的鸟苷酸激酶(guanylate kinase),命名为OsGK1。OsGK1在各器官中组成型表达,并在花后6 d的胚乳中表达水平最高。突变体胚乳中淀粉合成相关蛋白水平显著降低,尤其是AGPS2b和PHOI。此外,突变体fse2的胚发育严重受损,导致种子纯合致死;线粒体定位的AOX积累显著增强,而野生型中几乎检测不到,表明线粒体呼吸途径受损。【结论】由于OsGK1的功能缺陷,导致水稻种子中线粒体和造粉体发育异常,进而产生了胚致死以及胚乳粉质皱缩的表型,因此OsGK1对水稻种子的发育至关重要。
中图分类号:
李景芳, 田云录, 刘喜, 刘世家, 陈亮明, 江玲, 张文伟, 徐大勇, 王益华, 万建民. 鸟苷酸激酶OsGK1对水稻种子发育至关重要[J]. 中国水稻科学, 2018, 32(5): 415-426.
Jingfang LI, Yunlu TIAN, Xi LIU, Shijia LIU, Liangming CHEN, Ling JIANG, Wenwei ZHANG, Dayong XU, Yihua WANG, Jianmin WAN. The Guanylate Kinase OsGK1 is Essential for Seed Development in Rice[J]. Chinese Journal OF Rice Science, 2018, 32(5): 415-426.
图1 野生型与突变体fse2成熟种子表型比较 A,B,D—滇粳优1号(DJY)与fse2的成熟种子的外观表型;A,DJY(左),fse2(右)。B和D,DJY(上),fse2(下)。C—DJY(左)与fse2(右)种子的横切面。E,F—DJY(E)和fse2(F)成熟种子横切面的SEM观察。G~J—花后10 d I2-KI染色的DJY(G,H)和fse2(I,J)胚乳半薄切片。K—DJY和fse2的千粒重比较,n=3;L—DJY和fse2的粒长、粒宽和粒厚,n=20。所有数值为平均值±标准差;**野生型与突变体间的差异达0.01显著水平(t测验)。
Fig. 1. Phenotypic comparison of mature seeds of wild type and fse2 mutant. A, B, D, Mature seeds of Dianjingyou 1(DJY) and fse2. A, DJY (left), fse2 (right). B, D, DJY (upper), fse2 (lower). C, Cross-sections of mature seeds of DJY(left) and fse2 (right). E, F, Scanning electron microscopic (SEM) analysis of cross-sections of mature seeds of DJY (E) and fse2 (F). G-J, Semi-thin sections of DJY (G, H) and fse2 (I, J) endosperm at 10 days after pollination (DAP) stained with I2-KI. K, The 1000-grain weight of DJY and fse2. n=3. L, Grain length, width and thickness in DJY and fse2. n=20. All values are expressed as mean±SD. **means significant difference at 0.01 level (Student’s t-test).
图2 野生型与突变体fse2成熟种子理化特性分析 A~C—滇粳优1号(DJY)和fse2胚乳中总淀粉(A)、直链淀粉(B)和脂肪(C)含量的测定,n=3,取平均值±标准差,采用t测验,**P<0.01;D—DJY和fse2支链淀粉链长分布;E—DJY和fse2淀粉的RVA谱分析;F—DJY与fse2米粉的膨胀体积比较(n=3);G—DJY与fse2的尿素膨胀。
Fig. 2. Physicochemical characteristics of mature seeds of fse2 and its wild type. A~C, The contents of total starch (A), amylose (B), and lipid (C) in the endosperm of Dianjingyou(DJY) and fse2. n=3, Values are means±SD, Student’s t-test, **P<0.01. D, Amylopectin chain length distributions of DJY and fse2. E, Analysis of RVA characteristic of starch in DJY and fse2. F, The swollen volume of DJY and fse2 starch in urea solution (n=3). G, Starch expansion of DJY and fse2 in urea solutions.
试材 Test material | 最高黏度 Peak viscosity | 热浆黏度 Hot pasting viscosity | 崩解值Breakdown viscosity | 冷胶黏度 Cool pasting viscosity | 消减值 Setback viscosity | 峰值时间 Peak time / min | 糊化温度 Gelatinization temperature /℃ |
---|---|---|---|---|---|---|---|
DJY | 3197 | 1677 | 1520 | 3225 | 28 | 5.67 | 76.00 |
fse2 | 293 | 34 | 259 | 83 | -210 | 3.80 | 75.95 |
表1 野生型和突变体fse2淀粉的RVA谱特征分析
Table 1 Analysis of RVA characteristic values of starch in wild type and fse2 mutant.
试材 Test material | 最高黏度 Peak viscosity | 热浆黏度 Hot pasting viscosity | 崩解值Breakdown viscosity | 冷胶黏度 Cool pasting viscosity | 消减值 Setback viscosity | 峰值时间 Peak time / min | 糊化温度 Gelatinization temperature /℃ |
---|---|---|---|---|---|---|---|
DJY | 3197 | 1677 | 1520 | 3225 | 28 | 5.67 | 76.00 |
fse2 | 293 | 34 | 259 | 83 | -210 | 3.80 | 75.95 |
年份 Year | 透明种子数 Number of normal seeds | 粉质皱缩种子数 Number of floury and shrunken seeds | χ2(3:1) |
---|---|---|---|
2017 | 681 | 208 | 1.508 |
2016 | 573 | 172 | 1.612 |
表2 fse2遗传分析
Table 2 Genetic analysis of fse2.
年份 Year | 透明种子数 Number of normal seeds | 粉质皱缩种子数 Number of floury and shrunken seeds | χ2(3:1) |
---|---|---|---|
2017 | 681 | 208 | 1.508 |
2016 | 573 | 172 | 1.612 |
图3 FSE2的图位克隆 A―FSE2位点的图位克隆,FSE2定位在第3染色体短臂标记LJF3-16与LJF3-17之间约106 kb区域内,包含11个预测基因;B―Os03g0320900基因结构和突变位点,序列中存在3处单碱基替换(红色三角)和70 bp的缺失(红色方框)。引物F/R用于鉴定转基因家系CP-1、CP-2中透明种子的遗传背景;C―PCR鉴定FSE2基因中70 bp的缺失;D―FSE2蛋白的定位信号及结构域,在fse2中共有68个氨基酸,但只有N端20个氨基酸与野生型保持一致(虚线左边);mTP―线粒体转运肽;cTP―叶绿体转运肽;aa―氨基酸;E―FSE2在DJY和fse2发育种子中(开花10 d后)的相对表达量;F―具突变体背景的阳性转基因种子(CP-1 #3和CP-2 #7)恢复为正常表型,CP-1和CP-2为具有杂合突变体背景的转基因阳性家系,标尺为1 cm;G,H―CP-1家系(G)和CP-2家系(H)后代透明种子的PCR鉴定。G和H中上图均为转基因阳性鉴定结果,下图均为遗传背景检测结果,红色星号表示纯合突变体背景的阳性转基因种子。DJY―滇粳优1号。
Fig. 3. Map-based cloning of FSE2. A, Map-based cloning of the FSE2 locus. The FSE2 locus was mapped to a 106 kb region by markers LJF3-16 and LJF3-17 on the short arm of chromosome 3, which contains 11 predicted open reading frames (ORFs). B, The structure of Os03g0320900 and the mutation site. Three nucleotide substitutions (red triangles) and a 70 bp deletion (red box) in the sequence are indicated. The genetic background was identified with the primer pair F/R in transgenic lines. C, PCR analysis of the 70 bp deletion in the genomic region of Os03g0320900. D, Target peptides and functional domain of FSE2 protein. A total of 68 amino acids in fse2, and only the first 20 amino acids in the N-terminus are consistent with the wild type (left of the dotted line). mTP, Mitochondrial transit peptide; cTP, Chloroplast transit peptide; aa, amino acids. E, The relative expression levels of developing seeds (10 days post-pollination) in DJY and fse2. F, Positive transgenic seeds with homozygous fse2 background (CP-1 #3 and CP-2 #7) showed transparent endosperm. CP-1 and CP-2 are two positive transgenic lines with heterozygous background. Bar=1 cm. G-H, PCR analyses of the transparent seeds from CP-1 and CP-2 lines. The upper panels in G and H represent positive transgenic individuals. The lower panels of G and H show the backgrounds of these positive transgenic individuals. The red asterisks indicate the positive transgenic seeds with homozygous fse2 background. DJY, Dianjingyou 1.
图4 FSE2及其同源蛋白序列比对红色方框区域代表鸟苷酸激酶催化位点;GenBank蛋白质登录号:水稻,XP_051628708.1;拟南芥,NP_566276.1;玉米,NP_001149581.1;二穗短柄草,XP_003561683.1;毛果杨,XP_006380050.1;葡萄,XP_002279802.1;大豆,XP_003542897.2。
Fig. 4. Protein sequence alignment of FSE2 and its homologs. The red box indicates the catalytic site of GK; GenBank protein accession number are as follows O. sativa, XP_051628708.1; A. thaliana, NP_566276.1; Z. mays, NP_001149581.1; B. distachyon, XP_003561683.1; P. trichocarpa, XP_006380050.1; V. vinifera, XP_002279802.1; G. max, XP_003542897.2.
图5 FSE2的表达模式 A—FSE2在叶、穗子、鞘、根中的表达量;B—FSE2在花后3、6、9、12、15和18 d的发育胚乳中的表达量;C~F—根(C)、叶(D)、鞘(E)、小花(F)的GUS染色;G~J—开花后6(G)、9(H)、12(I)和15 d(J)的发育胚乳的GUS染色。
Fig. 5. The expression patterns of FSE2. A, Expression levels of FSE 2 in leaf, panicle, sheath and root of wild type. B, Expression levels of FSE2 in the developing endosperms of 3, 6, 9, 12, 15 and 18 days after flowering. C-F, GUS staining patterns in root (C), leaf (D), sheath (E) and panicle (F); G-J, GUS staining patterns in developing endosperm of 6 (G), 9 (H), 12 (I) and 15 days after flowering (J).
图6 突变体fse2的胚和胚乳发育异常 A—野生型和突变体fse2种子萌发后2 d的表型;B,C—野生型和突变体fse2种子吸胀后的胚(30℃下吸胀9 h),比例尺为1 mm;D,E—花后15 d野生型和突变体的胚,比例尺为1 mm;F—TTC法测定种子活力;G—Western blotting检测野生型和突变体成熟种子中的线粒体相关蛋白含量。CytC―细胞色素c合成酶C;COX2―细胞色素c氧化酶亚基2;NADH9―NADH脱氢酶亚基9;ATPβ―ATP合酶F0亚基6;AOX―交替氧化酶;H—Western blotting检测野生型和突变体成熟种子中淀粉合成相关酶蛋白含量,AGPL2―腺苷葡萄糖焦磷酸化酶大亚基2;AGPS2b―腺苷葡萄糖焦磷酸化酶小亚基2b;PPDKB―胞质丙酮酸磷酸双激酶B;PHOⅠ―质体磷酸化酶Ⅰ;BEⅠ―淀粉分支酶Ⅰ;BEⅡb―淀粉分支酶Ⅱb;SSⅡa―淀粉合酶Ⅱa;G,H中以ACTIN作为内参。DJY―滇粳优1号。
Fig. 6. The abnormal development of embryo and endosperm in the fse2 mutant. A, Comparison of the seed of wild type and fse2 after germinating for two days. B,C, Embryos of wild type and fse2 mutant seeds after imbibition (30℃, 9 h). Bars=1 mm. D, E, Developing embryos of 15 DAP seeds of wild type and fse2. Bars=1 mm. F, The determination of seed viability by TTC staining. G, Western blotting analysis of mitochondrial proteins in mature seeds of wild type and mutant. CytC, Cytochrome c biogenesis C; COX2, Cytochrome c oxidase subunit 2; NADH9, NADH dehydrogenase subunit 9; ATPβ, ATP synthase F0 subunit 6; AOX, Alternative oxidase. H, Western blotting analysis of starch synthesis enzymes in mature seeds of wild type and mutant. AGPL2, ADP-glucose pyrophosphorylase large subunit 2; AGPS2b, ADP-glucose pyrophosphorylase small subunit 2b; PPDKB, Cytosolic pyruvate orthophosphate dikinase B; PHOⅠ, Plastid phosphorylaseⅠ; BEⅠ, Starch-branching enzyme; BEⅡb, Starch-branching enzymeⅡb; SSⅡa, Starch synthaseⅡa. ACTIN antibody was used as a loading control in G and H. DJY, Dianjingyou 1.
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