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作物学报

作物学报 ›› 2022, Vol. 48 ›› Issue (2): 292-303.doi: 10.3724/SP.J.1006.2022.12013

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

水稻胚胎和胚乳双缺陷突变体eed1的表型与遗传分析

杨谨1,2(), 白爱宁3, 白雪3, 陈娟3, 郭林1, 刘春明1,3,*()   

  1. 1中国农业科学院作物科学研究所, 北京100081
    2中国农业科学院研究生院, 北京 100081
    3中国科学院植物研究所, 北京 100093
  • 收稿日期:2021-02-18 接受日期:2021-04-26 出版日期:2022-02-12 网络出版日期:2021-05-17
  • 通讯作者: 刘春明
  • 作者简介:E-mail: jinyang201587@163.com
  • 基金资助:
    本研究由北京市科技计划项目(Z181100002418010);中国科学院国际合作局国际伙伴计划(151111KYSB20180049);国家转基因生物新品种培育重大专项资助(2019ZX08010-001)

Phenotypic and genetic analyses of a rice mutant eed1 with defected embryo and endosperm development

YANG Jin1,2(), BAI Ai-Ning3, BAI Xue3, CHEN Juan3, GUO Lin1, LIU Chun-Ming1,3,*()   

  1. 1Institute of Crop Sciences, Chinese Academy of Agricultural Sciences, Beijing 100081, China
    2Graduate School, Chinese Academy of Agricultural Sciences, Beijing 100081, China
    3Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China
  • Received:2021-02-18 Accepted:2021-04-26 Published:2022-02-12 Published online:2021-05-17
  • Contact: LIU Chun-Ming
  • Supported by:
    This study was supported by the Beijing Scientific and Technological Research Program(Z181100002418010);the CAS-Commonwealth Scientific and Industrial Research Organization Bilateral Collaboration Project(151111KYSB20180049);the National Major Project for Developing New GM Crops(2019ZX08010-001)

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摘要：

从由化学诱变剂甲基磺酸乙酯(ethylmethane sulfonate, EMS)处理的粳稻品种中花11突变体库中, 筛选到一个可稳定遗传的胚胎和胚乳发育缺陷的突变体, 命名为embryo and endosperm defective 1 (eed1)。eed1籽粒千粒重、颖果的粒长、粒宽、粒厚、萌发率、总淀粉、直链淀粉和贮藏蛋白系列指标较野生型均显著降低。eed1颖果严重皱缩, 且胚乳呈粉质。利用扫描电镜观察发现, 与中花11相比, eed1胚乳中淀粉粒排列疏松, 多以单一、分散的淀粉粒存在, 且呈近似球形。eed1胚胎结构异常, 部分颖果未见有胚胎分化的痕迹。qRT-PCR发现, eed1胚乳中参与淀粉和贮藏蛋白合成的大部分基因表达下调。利用eed1与籼稻南京6号杂交得到的F2分离群体进行基因定位分析, 将EED1定位于9号染色体长臂672 kb的范围内, 包含114个开放读码框。本研究为进一步解析EED1基因调控水稻胚胎和胚乳发育的机制奠定了基础。

关键词: 水稻, 胚胎和胚乳发育, 籽粒灌浆

Abstract:

A stably inherited embryo and endosperm defective mutant, named embryo and endosperm defective 1 (eed1), was obtained from the mutant population of rice Zhonghua 11, generated by ethylmethane sulfonate (EMS). The 1000-grain weight, grain length, grain width, grain thickness, germination rate, total starch, amylose, and storage protein contents in eed1 mature caryopses were significantly decreased compared with wild type. In eed1, the caryopses were shrunken and the endosperm was opaque. Scanning electron microscopy observation revealed that starch granules in eed1 endosperm cells were loosely packed, in single, disperse and spherical forms. The embryo of eed1 was abnormal, and some caryopses showed no sign of embryonic differentiation. Most genes involved in biosynthesis of starch and storage protein were down-regulated in eed1 endosperm by qRT-PCR. EED1 was mapped in an interval of 672 kb on chromosome 9 using a F2 population derived from a cross between eed1 and Nanjing 6. The region contained 114 open reading frames. This study lays a foundation for further studying EED1 gene in regulating development of rice embryo and endosperm.

Key words: rice, embryo and endosperm development, grain filling

表1

实时荧光定量PCR分析所用的引物"

基因名
Genes ID		 正向引物序列
Forward sequences (5'-3')		 反向引物序列
Reverse sequences (5'-3')
Ubiquitin CTGTCAACTGCCGCAAGAAG GGCGAGTGACGCTCTAGTTC
OsAGPL1 GGAAGACGGATGATCGAGAAAG CACATGAGATGCACCAACGA
OsAGPL2 AGTTCGATTCAAGACGGATAGC CGACTTCCACAGGCAGCTTATT
OsAGPL3 AAGCCAGCCATGACCATTTG CACACGGTAGATTCACGAGACAA
OsAGPS1 GTGCCACTTAAAGGCACCATT CCCACATTTCAGACACGGTTT
OsAGPS2a ACTCCAAGAGCTCGCAGACC GCCTGTAGTTGGCACCCAGA
OsAGPS2b AACAATCGAAGCGCGAGAAA GCCTGTAGTTGGCACCCAGA
OsSS I GGGCCTTCATGGATCAACC CCGCTTCAAGCATCCTCATC
OsS IIa GCTTCCGGTTTGTGTGTTCA CTTAATACTCCCTCAACTCCACCAT
OsSS IIc GACCGAAATGCCTTTTTCTCG GGGCTTGGAGCCTCTCCTTA
OsSS IIIa GCCTGCCCTGGACTACATTG GCAAACATATGTACACGGTTCTGG
OsSS IIIb ATTCCGCTCGCAAGAACTGA CAACCGCAGGATAACGGAAA
OsSS IVa GGGAGCGGCTCAAACATAAA CCGTGCACTGACTGCAAAAT
OsSS IVb ATGCAGGAAGCCGAGATGTT ACGACAATGGGTGCCAAGAT
OsGBSS I
OsGBSS II		 AACGTGGCTGCTCCTTGAA
AGGCATCGAGGGTGAGGAG		 TTGGCAATAAGCCACACACA
CCATCTGGCCCACATCTCTA
OsBE I
OsBE IIa		 TGGCCATGGAAGAGTTGGC
GCCAATGCCAGGAAGATGA		 CAGAAGCAACTGCTCCACC
GCGCAACATAGGATGGGTTT
OsBE IIb ATGCTAGAGTTTGACCGC AGTGTGATGGATCCTGCC
OsISA1 TGCTCAGCTACTCCTCCATCATC AGGACCGCACAACTTCAACATA
OsISA2 TAGAGGTCCTCTTGGAGG AATCAGCTTCTGAGTCACCG
OsISA3 ACAGCTTGAGACACTGGGTTGAG GCATCAAGAGGACAACCATCTG
OsPUL ACCTTTCTTCCATGCTGG CAAAGGTCTGAAAGATGGG
OsDPE2 CAAGTACACCACAAGACCAGCAA CGTCCAACAGCGAATCCAAT
OsPHOL TTGGCAGGAAGGTTTCGCT CGAAGCCTGAAGTGAACTTGCT
OsPHOH CACCAAGACGAAGCTCATCAAG TTCACTCGTTGCTGGGTTCTC
GluA1 CATTTGAGCCAATTCGGAGT GGCCTGATTGTTGGAACTGT
GluA2 GCAAGAGCAGGAACAAGGAC CCTCATGGTGCAAAAGGTCT
GluA3 TGAAAACCAACCCTGACTCC ACTCATCTCCCCTGTTGTGC
GluB1 GCCAAAGTCAGAGCCAAAAG GAACCAATGTGCAACACCAG
GluB4 GCGACCAGAAGGCTACAAAG TTGCTTGTTGATCGTTGCTC
Glutelinλ TGGCGACCATAGCTTTCTCT GGGTTGTGCCATGGATTTAC
Globulin1 ATCGAGAACGGCGAGAAGT GGACGGAGATGGTATGGAGA
Globulin2 CGACGAGGTGTTCTACGTCA GTGTTGGCGGAGTAGACGAT
11S-Globulin CACCAAACCCGATCTTCAGT CGGAACAGCTTCTCCATCTC
19kD-Globulin GCCAGTAATTGCAGGGGATA AGGTCACCACCAACGTAAGC
Prolamin TCTCCAACCAACAATAGCAATG TGCGTAGCTATCTGTGCCCGTC
10kD-Prolamin TGCAGTATTTCCCACCAACA ACATGAACATGGCTGTGGAG
13kD-Prolamin CACAGCGCAGTTTGATGTTT GCTTGCCGCAATGCTATACT
17kD-Prolamin TTTGATGCTTGCACCTATGG GCAGCTGCTCAGTTTTAGCC
RA16 AGGTAGTGATCTCGGCGTTG CCGATTCCTGGCTGACATAG
RA17 TTCTCGGTATTGCTCCTCGT CTTATTCCTGGCCGACATTG

表2

EED1基因定位所用分子标记"

引物名
Primers ID		 正向引物序列
Forward sequences (5'-3')		 反向引物序列
Reverse sequences (5'-3')
OS907 CTTCTAACTTTTCCATCACATCG CCTCCAACTACCAGCCATAA
InDel950
InDel976		 ACTTTCTAACTTCCTAATAAG
GTTAAATCTTCTCTCAATCT		 GTTCTGGAATGAAATATTAGA
TCTCTCCGGTCTCTCCCTGC
RM107 AGATCGAAGCATCGCGCCCGAG ACTGCGTCCTCTGGGTTCCCGG
RM201 GTACTCTCGCCGTTCACAACTCC TTAGTGACCGGGATGACACAGC
RM245 ATGCCGCCAGTGAATAGC CTGAGAATCCAATTATCTGGGG
InDel908 TGGGAGAACTAAGGAGCAA CTGTTATCTGGCAATCTG

图1

ZH11与eed1成熟颖果表型分析 A: ZH11与eed1整粒成熟颖果外观比较, 标尺为1 mm; B: ZH11与eed1成熟颖果横切面比较, 标尺为1 mm; C: ZH11与eed1的成熟颖果大小比较, n = 20; D: ZH11与eed1的成熟烘干籽粒千粒重比较, n = 3。数值为平均值±标准差; 采用Student’s t测验, **表示P < 0.01; E~F: ZH11 (E)与eed1 (F)成熟颖果扫描电镜观察, 标尺为5 μm。"

"

图3

ZH11与eed1再生植株颖果表型分析 A~B: eed1愈伤组织在RE2培养基上分化产生不定芽和不定根(A), 最终长成小苗(B), 标尺为2 cm; C~D: ZH11与eed1不同发育时期颖果的鲜重(C)和干重(D)变化; E: ZH11与eed1受粉后0~30 d新鲜颖果的表型, 标尺为1 mm; F: ZH11与eed1受粉后24、27、30 d颖果烘干之后的表型, 标尺为1 mm; DAP为受粉后天数; 数值为平均值±标准差; 采用Student’s t测验, *表示P < 0.05, **表示P < 0.01。"

图4

eedl颖果中淀粉含量和贮藏蛋白累积分析 A~B: ZH11、WT和eed1成熟颖果的总淀粉(A)和直链淀粉(B)含量的测定, n = 3。数值为平均值±标准差; 采用Student’s t测验, **表示P < 0.01; C: ZH11、WT和eed1成熟颖果总蛋白SDS-PAGE电泳图谱; αGT: 谷蛋白酸性亚基, αGlb: 球蛋白; βGT: 谷蛋白碱性亚基; Pro or Alb: 醇溶蛋白或清蛋白。"

图5

eed1胚乳中淀粉和贮藏蛋白合成基因的表达水平分析 A: 参与淀粉生物合成的基因在ZH11与eed1胚乳中的相对表达水平; B: 参与贮藏蛋白生物合成的基因在ZH11与eed1胚乳中的相对表达水平。"

表3

EED1/eed1所产生籽粒的遗传分析"

表型
Phenotype		 杂合植株-1
Heterozygous plant-1		 杂合植株-2
Heterozygous plant-2		 杂合植株-3
Heterozygous plant-3
正常籽粒 Normal grains 157 152 133
胚胎且胚乳缺陷籽粒 eed1 grains 47 50 45
χ2(3:1) 0.418 < 3.84 (P > 0.05) 0.006 < 3.84 (P > 0.05) 0.007 < 3.84 (P > 0.05)

图6

EED1基因的遗传定位 A: EED1基因与9号染色体长臂上的分子标记OS907和InDel908连锁; B: 利用61个分离单株将EED1定位到672 kb的区间内, 该区间内有114个ORFs。"

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