中国水稻科学 ›› 2018, Vol. 32 ›› Issue (4): 325-334.DOI: 10.16819/j.1001-7216.2018.7122
收稿日期:
2017-09-28
修回日期:
2018-01-14
出版日期:
2018-07-10
发布日期:
2018-07-10
通讯作者:
李泽福
基金资助:
Kunneng ZHOU, Jiafa XIA, Tingchen MA, Yuanlei WANG, Zefu LI*()
Received:
2017-09-28
Revised:
2018-01-14
Online:
2018-07-10
Published:
2018-07-10
Contact:
Zefu LI
摘要: 【目的】对叶绿体发育相关基因进行克隆和功能分析,为解析叶绿体功能奠定分子基础。【方法】用甲基磺酸乙酯(EMS)处理籼稻9311获得一个条纹叶和白穗突变体slwp,通过色素分析和农艺性状观察分析该突变体的表型,通过图位克隆方法分离该基因,进一步利用定量PCR分析相关基因的表达情况。【结果】突变体slwp从2叶期开始至抽穗期表现出条纹叶表型,抽穗后幼穗白化,光合色素含量明显低于野生型;株高降低、抽穗延迟、产量降低等表型。该突变性状为单隐性核基因控制,该基因定位于水稻第6染色体短臂C6-4和N14标记之间0.91 Mb区间内。基因组测序表明核糖核苷二磷酸还原酶小亚基基因(RNRS1)编码区第776位点发生单碱基替换,导致甘氨酸突变为天冬氨酸;该基因与已报导的水稻基因St1、Gws和St-wp为等位基因。通过对这4个等位基因的突变位点和表型进行分析,总结了该基因不同位点突变对植株表型的影响以及籼粳之间的差异。表达分析显示与叶绿素合成有关的基因受到不同程度调控,叶绿体发育第一和第二阶段基因上调表达,光合作用相关基因均下调表达。【结论】本研究分析了SLWP(RNRS1)基因不同位点的变异对水稻表型的影响,相关结果加深了对RNRS1基因功能的认识,有助于阐明叶绿体发育的分子机制。
中图分类号:
周坤能, 夏加发, 马廷臣, 王元垒, 李泽福. 水稻条纹叶和白穗基因SLWP的定位及变异分析[J]. 中国水稻科学, 2018, 32(4): 325-334.
Kunneng ZHOU, Jiafa XIA, Tingchen MA, Yuanlei WANG, Zefu LI. Mapping and Mutation Analysis of Stripe Leaf and White Panicle Gene SLWP in Rice[J]. Chinese Journal OF Rice Science, 2018, 32(4): 325-334.
引物用途与名称 Usage and name | 正向引物 Forward sequence(5′-3′) | 反向引物 Reverse sequence(5′-3′) |
---|---|---|
定位和测序引物 Primers for mapping and sequencing | ||
C6-4 | CAGTTAACACCAATCCAATCCA | CCAAATGGGCAGTAGTTTGAA |
C6-5 | GCTTCTCCCGGAGTATGTCA | TGGTCTGAAAAGTGCCAAAA |
C6-6 | CCCCAAGCTTGTTTGTCTCT | TTGGGTCTTTGAGCTTCCTC |
N4 | TCAAGTTGCTAAACCTTATCTG | AGATGAACTGTGCTAAAAGATG |
N12 | GTAACTTAAAAGCCAATGTTGA | GAGTACTACCATCCATCCCTGT |
N14 | TCTGTGGACGTAGTAGGTTGA | CCTTCCTTAGGTCTGGCTC |
slwp-g | ACAACCCCAAATCCCCATCCA | ACCACGCGCATGTATTACTACT |
定量PCR引物 Primers for quantitative RT-PCR | ||
PORA | ATCACCAAGGGCTACGTCTC | GAGTTGTTGTTCCAGCTCCA |
HEMA1 | CACCAGTCTGAATCATAT | CTACCACTTCTCTAATCC |
YGL1 | TGGACAGTTGAAGATGTT | GAATAGGACGGTAAGGTT |
CHLI | AGTAACCTTGGTGCTGTG | AATCCATCAACATTCAACTCTG |
CHLH | CTATACATTCGCCACACT | TATCACACAACTCCCAAG |
CHLD | GGAAAGAGAGGGCATTAG | CAATACGATCAAGTAAGTGTT |
FtsZ | GTTGGTGTTTCTTCCAGCAA | CCTCAATAGACGACCCGATT |
RpoTp2 | AAGTCTGGCTTACGCTGGTT | AGGATCCTCAGCATTCATCC |
rpoA | AAATCGTTGATACGGCACAA | ATTCACATTTCGAACAGGCA |
rpoB | GCATTGTTGGAACTGGATTG | GCCGATGGGTAACTAAAGGA |
rbcL | GTTGAAAGGGATAAGTTGA | AATGGTTGTGAGTTTACG |
rbcS | TCATCAGCTTCATCGCCTAC | ACTGGGAACACACGAAACAA |
psbA | AAGTTTCTCTGATGGTATG | ATAGCACTGAATAGGGAA |
psaA | GAGATACCACTTCCTCAT | ACTAAGAAATTCTGCGTATT |
psaB | TTGGTATTGCTACCGCACAT | CCGGACGTCCATAGAAAGAT |
psbB | TCATATTGCTGCGGGTACAT | AGTTGCTGACCCATACCACA |
psbC | TACAACCTTGGCAAGAACGA | TACGCCACCCACAGAATTTA |
SLWP | AGTAATACATGCGCGTGGTG | ACACGGGCAGCTGATACTAA |
RNRS2 | AGATGTACAACGTCGCCAAC | GACATTACGGACGCCTTCTG |
Ubq | GCTCCGTGGCGGTATCAT | CGGCAGTTGACAGCCCTAG |
表1 本研究所用引物的序列
Table 1 Primer sequences used in the study.
引物用途与名称 Usage and name | 正向引物 Forward sequence(5′-3′) | 反向引物 Reverse sequence(5′-3′) |
---|---|---|
定位和测序引物 Primers for mapping and sequencing | ||
C6-4 | CAGTTAACACCAATCCAATCCA | CCAAATGGGCAGTAGTTTGAA |
C6-5 | GCTTCTCCCGGAGTATGTCA | TGGTCTGAAAAGTGCCAAAA |
C6-6 | CCCCAAGCTTGTTTGTCTCT | TTGGGTCTTTGAGCTTCCTC |
N4 | TCAAGTTGCTAAACCTTATCTG | AGATGAACTGTGCTAAAAGATG |
N12 | GTAACTTAAAAGCCAATGTTGA | GAGTACTACCATCCATCCCTGT |
N14 | TCTGTGGACGTAGTAGGTTGA | CCTTCCTTAGGTCTGGCTC |
slwp-g | ACAACCCCAAATCCCCATCCA | ACCACGCGCATGTATTACTACT |
定量PCR引物 Primers for quantitative RT-PCR | ||
PORA | ATCACCAAGGGCTACGTCTC | GAGTTGTTGTTCCAGCTCCA |
HEMA1 | CACCAGTCTGAATCATAT | CTACCACTTCTCTAATCC |
YGL1 | TGGACAGTTGAAGATGTT | GAATAGGACGGTAAGGTT |
CHLI | AGTAACCTTGGTGCTGTG | AATCCATCAACATTCAACTCTG |
CHLH | CTATACATTCGCCACACT | TATCACACAACTCCCAAG |
CHLD | GGAAAGAGAGGGCATTAG | CAATACGATCAAGTAAGTGTT |
FtsZ | GTTGGTGTTTCTTCCAGCAA | CCTCAATAGACGACCCGATT |
RpoTp2 | AAGTCTGGCTTACGCTGGTT | AGGATCCTCAGCATTCATCC |
rpoA | AAATCGTTGATACGGCACAA | ATTCACATTTCGAACAGGCA |
rpoB | GCATTGTTGGAACTGGATTG | GCCGATGGGTAACTAAAGGA |
rbcL | GTTGAAAGGGATAAGTTGA | AATGGTTGTGAGTTTACG |
rbcS | TCATCAGCTTCATCGCCTAC | ACTGGGAACACACGAAACAA |
psbA | AAGTTTCTCTGATGGTATG | ATAGCACTGAATAGGGAA |
psaA | GAGATACCACTTCCTCAT | ACTAAGAAATTCTGCGTATT |
psaB | TTGGTATTGCTACCGCACAT | CCGGACGTCCATAGAAAGAT |
psbB | TCATATTGCTGCGGGTACAT | AGTTGCTGACCCATACCACA |
psbC | TACAACCTTGGCAAGAACGA | TACGCCACCCACAGAATTTA |
SLWP | AGTAATACATGCGCGTGGTG | ACACGGGCAGCTGATACTAA |
RNRS2 | AGATGTACAACGTCGCCAAC | GACATTACGGACGCCTTCTG |
Ubq | GCTCCGTGGCGGTATCAT | CGGCAGTTGACAGCCCTAG |
图1 突变体slwp和野生型的表型性状 A—2叶期和3叶期表型;B和C—分蘖期叶片表型;D—孕穗期叶鞘表型;E和F—抽穗期幼穗表型;G—种子表型。
Fig. 1. Phenotypic characteristics of the slwp mutant and its wild type(WT). A, Phenotypes of two- and three-leaf stages; B and C, Leaf phenotype of tillering stage; D, Sheath phenotype of booting stage; E and F—Panicle phenotype of heading stage; G, Seed phenotype.
图2 突变体slwp和野生型色素含量测定 chla―叶绿素a;chlb―叶绿素b;a+b―总叶绿素;Car―胡萝卜素。
Fig. 2. Pigments determination of the slwp mutant and its wild type(WT). Chla, Chlorophyll a; chlb, Chlorophyll b; a+b, Total chlorophyll; Car―Carotene.
农艺性状 Agronomic trait | 野生型 Wild type | 突变体 swlp |
---|---|---|
株高Plant height / cm | 124.7±3.4 | 98.2±4.8** |
播始历期Days from sowing to heading / d | 98.4±1.2 | 108.3±1.0** |
穗长Panicle length / cm | 25.5±1.1 | 22.1±1.3** |
有效穗数Number of effective panicles per plant | 7.1±1.4 | 6.6±1.7 |
每穗粒数Grain number per panicle | 245.1±32.6 | 145.0±21.0** |
结实率Seed-setting rate / % | 88.2±1.3 | 87.9±1.6 |
千粒重1000-grain weight / g | 30.8±0.4 | 27.9±0.6** |
理论产量Theoretical yield per plot / kg | 10.74±0.23 | 5.34±0.35** |
实际产量Actual yield per plot / kg | 9.16±0.26 | 4.43±0.47** |
表2 野生型和突变体的农艺性状分析
Table 2 Agronomic traits of the wild type (WT) and slwp mutant.
农艺性状 Agronomic trait | 野生型 Wild type | 突变体 swlp |
---|---|---|
株高Plant height / cm | 124.7±3.4 | 98.2±4.8** |
播始历期Days from sowing to heading / d | 98.4±1.2 | 108.3±1.0** |
穗长Panicle length / cm | 25.5±1.1 | 22.1±1.3** |
有效穗数Number of effective panicles per plant | 7.1±1.4 | 6.6±1.7 |
每穗粒数Grain number per panicle | 245.1±32.6 | 145.0±21.0** |
结实率Seed-setting rate / % | 88.2±1.3 | 87.9±1.6 |
千粒重1000-grain weight / g | 30.8±0.4 | 27.9±0.6** |
理论产量Theoretical yield per plot / kg | 10.74±0.23 | 5.34±0.35** |
实际产量Actual yield per plot / kg | 9.16±0.26 | 4.43±0.47** |
图3 SLWP基因的图位克隆 A—SLWP基因初定位;B—SLWP基因精细定位;C—SLWP基因结构和突变位点;ATG和TGA分别代表起始密码子和终止密码子。
Fig. 3. Map-based cloning of the SLWP gene. A, Initial mapping of the SLWP gene; B, Fine mapping of the SLWP gene; C, Structure and mutant site of the SLWP gene; ATG and TGA indicate initiation codon and termination codon, respectively.
图4 SLWP及其同源蛋白的序列比对序列来源于水稻LOC_Os06g14620(SLWP)、拟南芥At3g23580、大豆XP_003547645.1、玉米NP_001130908.2、烟草NP_001312237.1和小立碗藓XP_001768501.1;下划线代表核糖核苷酸还原酶结构域;ST1、STWP、GWS和SLWP蛋白突变位点以红色字体和线框标出。
Fig. 4. Alignment of SLWP and SLWP-related proteins of different organisms. Sequences are for Oryza sativa LOC_Os06g14620(SLWP), Arabidopsis thaliana At3g23580, Glycine max XP_003547645.1, Zea mays NP_001130908.2, Nicotiana tabacum NP_001312237.1 and Physcomitrella patens XP_001768501.1; Underline represents nucleotide reductase domain; Red font and frame indicate mutant sites of ST1, STWP, GWS and SLWP.
图5 蛋白三维结构预测 A—st1、st-wp和slwp;B—突变蛋白gws;红色表示突变位点。
Fig. 5. Prediction of 3D structure model. A—st1, st-wp and slwp; B—Mutant protein gws; The red regions represent mutant sites.
突变体 Mutant | 野生型 Wild type | 氨基酸突变位点 Mutant site of amino acid | 叶片表型 Leaf phenotype | 穗部表型 Panicle phenotype | 主要农艺性状变化 Agronomic trait change compared with its wild type |
---|---|---|---|---|---|
st1 | FL176 | p. Lys40Arg | 4或5叶期至分蘖期出现条纹;抽穗后表型恢复Display stripe from 4- or 5-leaf stage to tillering, then revert to normal after heading | 无 None | 无 None |
gws | 日本晴 Nipponbare | 104原有+插入14个氨基酸 104 original+Inserted 14 amino acid residues | 苗期至抽穗期叶片均出现条纹Display stripe from the seedling to heading stage | 无 None | 株高、穗长、有效分蘖数、结实率、千粒重、单株生物量均降低Decreased plant height, panicle length, tiller number, seed-setting rate, 1000-grain weight, and plant biomass per plant |
st-wp | 9311 | p. Glu103Val | 苗期至抽穗期叶片均出现条纹Display stripe from the seedling to heading stage | 白穗 White panicle | 株高、穗长、每穗粒数、结实率、千粒重降低,抽穗期延长,有效穗数不变 Decreased plant height, panicle length, grain number per panicle, seed-setting rate, and 1000-grain weight; prolonged heading date; similar tiller number |
slwp | 9311 | p. Gly259Asp | 苗期至抽穗期叶片出现条纹 Display stripe from the seedling to heading stage | 白穗 White panicle | 株高、穗长、每穗粒数、千粒重、理论产量、实际产量下降,抽穗期延长,有效穗数和结实率不变 Decreased plant height, panicle length, grain number per panicle, 1000-grain weight, theoretical yield, and actual yield; prolonged heading date, similar tiller number and seed-setting rate |
表3 SLWP变异分析
Table 3 Mutation analysis of SLWP.
突变体 Mutant | 野生型 Wild type | 氨基酸突变位点 Mutant site of amino acid | 叶片表型 Leaf phenotype | 穗部表型 Panicle phenotype | 主要农艺性状变化 Agronomic trait change compared with its wild type |
---|---|---|---|---|---|
st1 | FL176 | p. Lys40Arg | 4或5叶期至分蘖期出现条纹;抽穗后表型恢复Display stripe from 4- or 5-leaf stage to tillering, then revert to normal after heading | 无 None | 无 None |
gws | 日本晴 Nipponbare | 104原有+插入14个氨基酸 104 original+Inserted 14 amino acid residues | 苗期至抽穗期叶片均出现条纹Display stripe from the seedling to heading stage | 无 None | 株高、穗长、有效分蘖数、结实率、千粒重、单株生物量均降低Decreased plant height, panicle length, tiller number, seed-setting rate, 1000-grain weight, and plant biomass per plant |
st-wp | 9311 | p. Glu103Val | 苗期至抽穗期叶片均出现条纹Display stripe from the seedling to heading stage | 白穗 White panicle | 株高、穗长、每穗粒数、结实率、千粒重降低,抽穗期延长,有效穗数不变 Decreased plant height, panicle length, grain number per panicle, seed-setting rate, and 1000-grain weight; prolonged heading date; similar tiller number |
slwp | 9311 | p. Gly259Asp | 苗期至抽穗期叶片出现条纹 Display stripe from the seedling to heading stage | 白穗 White panicle | 株高、穗长、每穗粒数、千粒重、理论产量、实际产量下降,抽穗期延长,有效穗数和结实率不变 Decreased plant height, panicle length, grain number per panicle, 1000-grain weight, theoretical yield, and actual yield; prolonged heading date, similar tiller number and seed-setting rate |
图6 野生型和突变体中叶绿素合成、叶绿体发育和光合作用相关基因的表达分析 A—叶绿素合成相关基因表达分析;B—叶绿体发育相关基因表达分析;C—光合作用相关基因表达分析。*和**分别表示5%和1%显著水平。
Fig. 6. Expression analysis of genes involved in chlorophyll biosynthesis, chloroplast development and photosynthesis between wild type(WT) and mutant. A, Expression analysis of genes associated with chlorophyll biosynthesis; B, Expression analysis of genes associated with chloroplast development; C, Expression analysis of genes associated with photosynthesis. * and ** represent significant difference at 5% and 1% levels, respectively.
图7 SLWP(RNRS1)和RNRS2在中花11(ZH11)、9311和slwp突变体不同组织中的表达分析
Fig. 7. Expression analysis of SLWP (RNRS1) and RNRS2 in different tissues of ZH11, 9311 and slwp mutant.
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