中国水稻科学 ›› 2023, Vol. 37 ›› Issue (5): 497-506.DOI: 10.16819/j.1001-7216.2023.221201
徐欢1,#, 周涛2,#, 孙悦1, 王木妹3, 杨亚春2, 马卉2, 李浩2, 徐大伟1, 周海3, 杨剑波2, 倪金龙2,*()
收稿日期:
2022-12-05
修回日期:
2023-01-31
出版日期:
2023-09-10
发布日期:
2023-09-13
通讯作者:
*email: 作者简介:
第一联系人:#共同第一作者
基金资助:
XU Huan1,#, ZHOU Tao2,#, SUN Yue1, WANG Mumei3, YANG Yachun2, MA Hui2, LI Hao2, XU Dawei1, ZHOU Hai3, YANG Jianbo2, NI Jinlong2,*()
Received:
2022-12-05
Revised:
2023-01-31
Online:
2023-09-10
Published:
2023-09-13
Contact:
*email: About author:
First author contact:#These authors contributed equally to this work
摘要:
【目的】 鉴定水稻颖壳类病斑突变体,并进行基因定位,为基因克隆及其分子机制研究奠定基础。【方法】 对野生型材料LR005和经EMS诱变得到的颖壳类病斑突变体glmm1(glume lesion mimics mutant 1)进行农艺性状分析、扫描电镜分析、DAB染色和全硅含量测定。glmm1与广亲和材料L422杂交获得的F2群体用于遗传分析,利用图位克隆和BSA-seq方法进行基因定位。【结果】 突变体glmm1在抽穗10 d后颖壳和叶片逐渐出现褐色斑点,成熟后颖壳完全呈现褐色。与野生型相比,突变体的株高、穗长、每穗总粒数、结实率和千粒重等都极显著降低。DAB染色表明glmm1颖壳和叶片的活性氧含量增多;扫描电镜显示突变体颖壳和叶片表面硅质细胞皱缩。遗传分析结果表明,突变体glmm1的颖壳类病斑表型受到一对隐性基因控制。利用glmm1与L422的F2分离群体,通过图位克隆和BSA-seq等策略将glmm1定位在水稻第2染色体上68 kb的区间内。该区间内有10个候选基因。序列分析发现该区间仅有一个SNP位点,位于基因Lsi1(LOC_Os02g51110)的第5个外显子上,导致第238位氨基酸由异亮氨酸变为苏氨酸。对颖壳和叶片全硅含量的测定表明,glmm1突变体中硅的积累减少,说明GLMM1可能是Lsi1的等位突变。【结论】 GLMM1是Lsi1新的等位突变基因,该突变造成植株硅含量的降低和活性氧的积累,致使颖壳和叶片产生褐色类病斑。
徐欢, 周涛, 孙悦, 王木妹, 杨亚春, 马卉, 李浩, 徐大伟, 周海, 杨剑波, 倪金龙. 水稻颖壳类病斑突变体glmm1的鉴定与基因定位[J]. 中国水稻科学, 2023, 37(5): 497-506.
XU Huan, ZHOU Tao, SUN Yue, WANG Mumei, YANG Yachun, MA Hui, LI Hao, XU Dawei, ZHOU Hai, YANG Jianbo, NI Jinlong. Characterization and Gene Mapping of a Glume Lesion Mimic Mutant glmm1 in Rice[J]. Chinese Journal OF Rice Science, 2023, 37(5): 497-506.
引物名称 Primer name | 正向引物 Forward primer (5'→3') | 反向引物 Reverse primer (5'→3') |
---|---|---|
X1 | ACAACTGACCTACTCACCTAC | CAATGCAGGTGACATATTTTCAA |
X2 | TGCCTCATACAAATGAGAGAC | ATAAAAGTAGGACGGCTCTTC |
X3 | ACAGGTCACGAAGTACTGAA | GGGCTATTTATGTGTGCCTT |
X4 | CCTTATCTTGGTTTGGTCTTTT | CATCTGCCATCCTACCTGAA |
X5 | TTTTGACCGAGGGAGTACG | TCAGGTGACCCCACGACT |
X6 | AGAGGTGGCTTCGCTATTG | TTTGGGACGGAGGGAGTA |
Y1 | GGGCCGTCTGTCTCTAAA | CACGAAAAACCTTGACCA |
Y2 | AGGCCCAAGTCATTCAAC | CGCAAAGCATGAGACAAT |
Y3 | CCTACGTAAGAAGCCATCG | TCGGTTTAGAGGGGATTG |
Y4 | AGGCTCCATCAAACCAAC | ATTTCACATGCCCCTCAT |
Y5 | CGCTCACACAAGTTAATTGGA | GCGTTAAAACGTATGGACAG |
Y6 | GTAACTGGGAGTACAGTGGTAA | AAATTAAAGCGACAGAGATTTCC |
表1 InDel标记引物信息
Table 1. Primer sequences of InDel markers for fine mapping of GLMM1.
引物名称 Primer name | 正向引物 Forward primer (5'→3') | 反向引物 Reverse primer (5'→3') |
---|---|---|
X1 | ACAACTGACCTACTCACCTAC | CAATGCAGGTGACATATTTTCAA |
X2 | TGCCTCATACAAATGAGAGAC | ATAAAAGTAGGACGGCTCTTC |
X3 | ACAGGTCACGAAGTACTGAA | GGGCTATTTATGTGTGCCTT |
X4 | CCTTATCTTGGTTTGGTCTTTT | CATCTGCCATCCTACCTGAA |
X5 | TTTTGACCGAGGGAGTACG | TCAGGTGACCCCACGACT |
X6 | AGAGGTGGCTTCGCTATTG | TTTGGGACGGAGGGAGTA |
Y1 | GGGCCGTCTGTCTCTAAA | CACGAAAAACCTTGACCA |
Y2 | AGGCCCAAGTCATTCAAC | CGCAAAGCATGAGACAAT |
Y3 | CCTACGTAAGAAGCCATCG | TCGGTTTAGAGGGGATTG |
Y4 | AGGCTCCATCAAACCAAC | ATTTCACATGCCCCTCAT |
Y5 | CGCTCACACAAGTTAATTGGA | GCGTTAAAACGTATGGACAG |
Y6 | GTAACTGGGAGTACAGTGGTAA | AAATTAAAGCGACAGAGATTTCC |
图1 野生型和突变体glmm1的表型比较 A, B−苗期和成熟期的野生型(WT)和glmm1的表型;C, D, E−WT和glmm1的籽粒;F, I−抽穗初期;G, J−抽穗中期;H, K−成熟期。标尺分别为15 cm(A, B), 1 cm(C, D, E)和4 cm(F, G, H, I, J, K)。
Fig. 1. Phenotypic comparison of WT and mutant glmm1. A and B, Phenotype of WT and glmm1 at seedling stage and maturity stage; C, D and E, Seeds of WT and glmm1; F and I, Early heading stage; G and J, Middle heading stage; H and K, Maturity stage. Bars, 15 cm (A, B), 1 cm (C, D, E), 4 cm (F, G, H, I, J, K), respectively.
材料 Material | 株高 Plant height / cm | 穗长 Panicle length/cm | 有效穗数 No. of effective panicles | 每穗总粒数 No. of spikelets per panicle | 结实率 Seed-setting rate / % | 千粒重 1000-grain weight / g |
---|---|---|---|---|---|---|
野生型WT | 92.06±4.24 | 20.59±1.43 | 5.6±2.2 | 535.8±62.9 | 42.07±8.41 | 20.23±0.14 |
glmm1 | 85.45±3.02** | 18.86±1.41** | 5.2±1.2 | 408.2±72.4** | 29.21±9.50** | 16.84±0.27** |
表2 野生型与突变体glmm1的主要农艺性状比较
Table 2. Agronomic traits of the wild type (WT) and glmm1.
材料 Material | 株高 Plant height / cm | 穗长 Panicle length/cm | 有效穗数 No. of effective panicles | 每穗总粒数 No. of spikelets per panicle | 结实率 Seed-setting rate / % | 千粒重 1000-grain weight / g |
---|---|---|---|---|---|---|
野生型WT | 92.06±4.24 | 20.59±1.43 | 5.6±2.2 | 535.8±62.9 | 42.07±8.41 | 20.23±0.14 |
glmm1 | 85.45±3.02** | 18.86±1.41** | 5.2±1.2 | 408.2±72.4** | 29.21±9.50** | 16.84±0.27** |
杂交组合 Cross | 野生型 Wild type | 突变体 Mutant | 分离比 Segregation ratio | χ23:1 |
---|---|---|---|---|
glmm1/L422 | 1499 | 501 | 2.99:1 | 0.003 |
表3 glmm1颖壳类病斑性状的遗传分析
Table 3. Genetic analysis of the lesion mimic trait on glumes of glmm1.
杂交组合 Cross | 野生型 Wild type | 突变体 Mutant | 分离比 Segregation ratio | χ23:1 |
---|---|---|---|---|
glmm1/L422 | 1499 | 501 | 2.99:1 | 0.003 |
基因注释号 | 功能注释 | 基因符号 |
---|---|---|
基因注释号 Gene annotation | 功能注释 Function note | 基因名称 Gene name |
LOC_Os02g51100 | 砷泵驱动ATP酶 Arsenical pump-driving ATPase | |
LOC_Os02g51110 | 水通道蛋白 Aquaporin protein | Lsi1 |
LOC_Os02g51119 | 表达蛋白 Expressed protein | |
LOC_Os02g51130 | 含有糖基转移酶8结构域的蛋白质 Glycosyl transferase 8 domain containing protein | |
LOC_Os02g51140 | 富氮蛋白 N-rich protein | |
LOC_Os02g51150 | 鞘脂C4-羟化酶SUR2 Sphingolipid C4-hydroxylase SUR2 | |
LOC_Os02g51160 | 水解酶,作用于C-N Hydrolase, acting on carbon-nitrogen | |
LOC_Os02g51164 | 表达蛋白Expressed protein | |
LOC_Os02g51170 | 表达蛋白Expressed protein | |
LOC_Os02g51180 | Cullin蛋白 Cullin protein | OsCUL3a |
表4 候选区间基因注释
Table 4. Gene annotation of candidate region
基因注释号 | 功能注释 | 基因符号 |
---|---|---|
基因注释号 Gene annotation | 功能注释 Function note | 基因名称 Gene name |
LOC_Os02g51100 | 砷泵驱动ATP酶 Arsenical pump-driving ATPase | |
LOC_Os02g51110 | 水通道蛋白 Aquaporin protein | Lsi1 |
LOC_Os02g51119 | 表达蛋白 Expressed protein | |
LOC_Os02g51130 | 含有糖基转移酶8结构域的蛋白质 Glycosyl transferase 8 domain containing protein | |
LOC_Os02g51140 | 富氮蛋白 N-rich protein | |
LOC_Os02g51150 | 鞘脂C4-羟化酶SUR2 Sphingolipid C4-hydroxylase SUR2 | |
LOC_Os02g51160 | 水解酶,作用于C-N Hydrolase, acting on carbon-nitrogen | |
LOC_Os02g51164 | 表达蛋白Expressed protein | |
LOC_Os02g51170 | 表达蛋白Expressed protein | |
LOC_Os02g51180 | Cullin蛋白 Cullin protein | OsCUL3a |
图4 水稻颖壳和叶片的扫描电镜观察 A, C, I, K−抽穗初期野生型(WT)和glmm1的穗和叶片的表型特征;B, D, J, L−抽穗初期WT和glmm1的穗和叶片的扫描电镜观察;E, G, M, O−成熟期WT和glmm1的穗和叶片的表型特征;F, H, N, P−成熟期WT和glmm1的穗和叶片的扫描电镜观察。标尺为4 cm(A, C, E, G, I, K, M, O),40 μm(B, D, F, H, J, L, N, P)。红色箭头指向植硅石和二氧化硅细胞。
Fig. 4. Scanning electron microscopy observation of rice glumes and leaves. A, C, I and K, Phenotypic characteristics of panicles and leaves of WT and glmm1 at the early heading stage; B, D, J and L, SEM of panicles and leaves of WT and glmm1 at the early heading stage; E, G, M and O, Phenotypic characteristics of panicles and leaves of WT and glmm1 at the ripening stage, respectively;F, H, N and P, SEM of panicles and leaves of WT and glmm1 at the ripening stage. Bars: 4 cm (A, C, E, G, I, K, M, O),40 μm (B, D, F, H, J, L, N, P). The red arrow indicates phytolith and silica cells.
图5 野生型和突变体glmm1的全硅含量测定 A−颖壳;B−叶片。**代表野生型与突变体间的差异达0.01显著水平(n=3)。
Fig. 5. Determination of total silicon content of the wild type (WT) and its mutant glmm1. A, Glume; B, Leaf. ** mean difference between WT and glmm1 is significant at 0.01 level (t-test, n=3).
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