中国水稻科学 ›› 2021, Vol. 35 ›› Issue (6): 535-542.DOI: 10.16819/j.1001-7216.2021. 210205
吴先美1,#, 李三峰1,#, 胡萍1, 何瑞1, 焦然2, 毛一剑1, 鲁草林1, 胡娟2, 林晗2, 吴荣梁1, 朱旭东1, 饶玉春2,*(), 王跃星1,*()
收稿日期:
2021-02-16
修回日期:
2021-03-23
出版日期:
2021-11-10
发布日期:
2021-11-10
通讯作者:
饶玉春,王跃星
作者简介:
#共同第一作者
基金资助:
Xianmei WU1,#, Sanfeng LI1,#, Ping HU1, Rui HE1, Ran JIAO2, Yijian MAO1, Caolin LU1, Juan HU2, Han LIN2, Rongliang WU1, Xudong ZHU1, Yuchun RAO2,*(), Yuexing WANG1,*()
Received:
2021-02-16
Revised:
2021-03-23
Online:
2021-11-10
Published:
2021-11-10
Contact:
Yuchun RAO, Yuexing WANG
About author:
#These authors contributed equally to this work
摘要:
【目的】克隆水稻分蘖相关基因,为构建理想株型水稻,提高粮食产量提供理论基础及有利基因资源。【方法】在常规大田种植条件下分别比较突变体htd3(high-tillering dwarf 3)与野生型在幼苗期、抽穗期和成熟期表型及主要农艺性状差异,利用图位克隆方法克隆候选基因,利用荧光定量PCR方法分析HTD3及独脚金内酯和脱落酸相关基因的表达水平,测序比对分析HTD3在147份种质资源中的自然变异情况。【结果】与野生型相比,突变体htd3的分蘖芽生长较快,分蘖数和有效穗数显著增多,株高、一次枝梗数和每穗粒数显著降低,结实率和千粒重没有显著变化。遗传分析表明,htd3多分蘖的性状受一对隐性核基因控制。图位克隆将HTD3基因定位在第12染色体CM8和CM10之间约63.5 kb的物理区间内,互补转基因实验证明该区间内LOC_Os12g21710为控制突变体多分蘖表型的基因。HTD3在野生型和突变体中呈组成型表达,该基因突变会引起部分独脚金内酯和脱落酸相关基因的表达水平上调。水稻品种中HTD3编码区G2674A自然变异使得分蘖数显著增多。【结论】HTD3是最近报道的T20/MIT1基因的一个新的等位基因,HTD3突变导致水稻出现分蘖适度增加,株高略矮的表型,在培育理想株型水稻和高产育种上具有较大的应用潜力。
吴先美, 李三峰, 胡萍, 何瑞, 焦然, 毛一剑, 鲁草林, 胡娟, 林晗, 吴荣梁, 朱旭东, 饶玉春, 王跃星. 水稻分蘖调控基因HTD3的克隆与功能研究[J]. 中国水稻科学, 2021, 35(6): 535-542.
Xianmei WU, Sanfeng LI, Ping HU, Rui HE, Ran JIAO, Yijian MAO, Caolin LU, Juan HU, Han LIN, Rongliang WU, Xudong ZHU, Yuchun RAO, Yuexing WANG. Cloning and Functional Analysis of Rice Tillering Regulatory Gene HTD3[J]. Chinese Journal OF Rice Science, 2021, 35(6): 535-542.
标记 Marker | 正向引物 Forward primer (5′→3′) | 反向引物 Reverse primer (5′→3′) |
---|---|---|
B12-6 | TGAAGCGGTTTGACTTTGACC | GGGGTGAAAACTGGTAGGGT |
B12-9 | TCCTTCTCGTTTATGAACTTATGG | TAGAGCAAAGCAGAGCCCAG |
M9 | TAATCCCCTGCACTCCATCC | GTGAACAACCAGCCGAGAAT |
M17 | GACTTTCTAGCATTGCCCACA | GCATTAACTGGGGCCATTGT |
M25 | GAGACGGCCAGCTTAGGTAG | CAGTGCTACAGAAACAGGGC |
M29 | CCGAACTCCAGTTTGTGAGG | CGGAAATCTGACGCTGGTAT |
M35 | AAAAGAACAACACAGCCCCT | ACAAGGAGCAATCTGGACCA |
CM5 | TTGTGAACAAGAGCCAACGG | CGCTGTTGGGCATTCTTTAAAG |
CM8 | GTGTGATCCATGGGTAGCCT | CAGAGCCCTATTAGTCTATTGCT |
CM10 | TCACATGATACCTCGCGAGT | CAGACGATTCTACACAACAGGA |
CM13 | AGCAGCCAAGATTAAGGAGGA | CGTCAGAGTGATTAGCAAAAGGA |
表1 HTD3精细定位所用分子标记
Table 1 Sequences of markers for HTD3 mapping.
标记 Marker | 正向引物 Forward primer (5′→3′) | 反向引物 Reverse primer (5′→3′) |
---|---|---|
B12-6 | TGAAGCGGTTTGACTTTGACC | GGGGTGAAAACTGGTAGGGT |
B12-9 | TCCTTCTCGTTTATGAACTTATGG | TAGAGCAAAGCAGAGCCCAG |
M9 | TAATCCCCTGCACTCCATCC | GTGAACAACCAGCCGAGAAT |
M17 | GACTTTCTAGCATTGCCCACA | GCATTAACTGGGGCCATTGT |
M25 | GAGACGGCCAGCTTAGGTAG | CAGTGCTACAGAAACAGGGC |
M29 | CCGAACTCCAGTTTGTGAGG | CGGAAATCTGACGCTGGTAT |
M35 | AAAAGAACAACACAGCCCCT | ACAAGGAGCAATCTGGACCA |
CM5 | TTGTGAACAAGAGCCAACGG | CGCTGTTGGGCATTCTTTAAAG |
CM8 | GTGTGATCCATGGGTAGCCT | CAGAGCCCTATTAGTCTATTGCT |
CM10 | TCACATGATACCTCGCGAGT | CAGACGATTCTACACAACAGGA |
CM13 | AGCAGCCAAGATTAAGGAGGA | CGTCAGAGTGATTAGCAAAAGGA |
引物名称 Primer name | 引物序列 Primer sequences (5′→3′) |
---|---|
HTD3-CPT-EcoRⅠ-F | ACGAATTCGAGCTCGGTACCTGGTTCTGTGACTAAAGCGC |
HTD3-CPT-HindⅢ-R | GGCCAGTGCCAAGCTTTCTCCGGGGCCCTGAATATTCCTCT |
表2 扩增HTD3完整表达单元的引物
Table 2 Primers for amplifying the complete expression unit of HTD3.
引物名称 Primer name | 引物序列 Primer sequences (5′→3′) |
---|---|
HTD3-CPT-EcoRⅠ-F | ACGAATTCGAGCTCGGTACCTGGTTCTGTGACTAAAGCGC |
HTD3-CPT-HindⅢ-R | GGCCAGTGCCAAGCTTTCTCCGGGGCCCTGAATATTCCTCT |
图1 野生型和htd3的形态性状比较 A~C-野生型和htd3在苗期(A),分蘖期(B)和成熟期(C)的植株表型,标尺为6 cm;D,E-野生型和htd3的分蘖数(D)和株高(E)随播种后日期的动态变化。数据为平均值±标准差(n=15);采用t测验,**表示在0.01水平上差异显著,*表示在0.05水平上差异显著。
Fig. 1. Morphological characterization of WT and htd3. A~C, Plant phenotypes of wild-type (WT) and htd3 in the seedling stage (A), tillering stage (B) , and mature stage (C) , bar= 6 cm; D~E, Tiller number (D) and plant height (E) of WT and htd3 after sowing. Values are means ± SD (n=15), t-test, *, ** indicate significant difference at P<0.05 and P<0.01, respectively.
农艺性状Agronomic trait | 野生型WT | 突变体htd3 |
---|---|---|
株高Plant height /cm | 97.08±2.78 | 83.76±1.54* |
有效穗数Effective panicles | 11.0±1.0 | 28.0±2.0** |
穗长Panicle length/cm | 21.52±0.98 | 20.82±0.62 |
一次枝梗数 Primary branches | 12.0±2.0 | 9.0±1.0* |
二次枝梗数Secondary branches | 26.0±3.0 | 23.0±4.0 |
每穗粒数Grains per panicle | 121.0±11.0 | 104.0±16.0* |
每株总粒数Total grains per plant | 1331±28 | 2018±36** |
千粒重1000-grain weight/g | 23.7±2.1 | 23.9±1.3 |
结实率Seed setting rate/% | 87.2±5.8 | 90.0±2.4 |
表3 野生型和htd3的主要农艺性状比较
Table 3 Comparison of agronomic traits between WT and htd3.
农艺性状Agronomic trait | 野生型WT | 突变体htd3 |
---|---|---|
株高Plant height /cm | 97.08±2.78 | 83.76±1.54* |
有效穗数Effective panicles | 11.0±1.0 | 28.0±2.0** |
穗长Panicle length/cm | 21.52±0.98 | 20.82±0.62 |
一次枝梗数 Primary branches | 12.0±2.0 | 9.0±1.0* |
二次枝梗数Secondary branches | 26.0±3.0 | 23.0±4.0 |
每穗粒数Grains per panicle | 121.0±11.0 | 104.0±16.0* |
每株总粒数Total grains per plant | 1331±28 | 2018±36** |
千粒重1000-grain weight/g | 23.7±2.1 | 23.9±1.3 |
结实率Seed setting rate/% | 87.2±5.8 | 90.0±2.4 |
图2 野生型和htd3分蘖芽表型观察 A~F-野生型和突变体htd3在5叶期、6叶期、7叶期的分蘖芽表型, A~C为野生型, D~F为突变体, 标尺为1 cm;G-野生型和突变体htd3在各叶期的分蘖长度, n/0中n表示叶位, 0表示主茎, 即1/0表示主茎第一片叶对应的分蘖芽, 2/0表示主茎第2片叶对应的分蘖芽, 以此类推。
Fig. 2. Phenotypic observation of tillering buds of WT and htd3. A~F, Phenotype of tillering buds of WT and htd3 at five-, six- and seven-leaf stage; A-C represents WT; D-F represents htd3; bars=1 cm; G, Tillering bud length of WT and htd3 at each leaf stage, in n/0, n represents the leaf position, 0 represents the main stem, 1/0 represents the tiller bud corresponding to the first leaf of the main stem, 2/0 represents the tiller bud corresponding to the second leaf of the main stem, and so on.
组合Cross | F1表型 Phenotype of F1 plants | F2群体 F2 population | χ2 (3:1) | χ20. 05 | ||
---|---|---|---|---|---|---|
总株数 No. of plants | 正常分蘖株数 No. of normal plants | 多分蘖株数 No. of high-tillering plants | ||||
中花11/ TN1 Zhonghua 11/TN1 | 正常分蘖Normal tiller | 864 | 666 | 198 | 2 | 3. 84 |
表4 水稻矮秆多分蘖突变体htd3的遗传分析
Table 4 Genetic analysis of rice high-tillering mutant htd3.
组合Cross | F1表型 Phenotype of F1 plants | F2群体 F2 population | χ2 (3:1) | χ20. 05 | ||
---|---|---|---|---|---|---|
总株数 No. of plants | 正常分蘖株数 No. of normal plants | 多分蘖株数 No. of high-tillering plants | ||||
中花11/ TN1 Zhonghua 11/TN1 | 正常分蘖Normal tiller | 864 | 666 | 198 | 2 | 3. 84 |
图3 HTD3的图位克隆和互补转基因验证 A-HTD3的初定位;B-HTD3的精细定位;C-定位区间内预测的ORF;D-候选基因LOC_Os12g21710的基因结构及在野生型和htd3中的序列差异,黑色方框代表外显子,白色方框代表UTR,黑色线条代表内含子;E-野生型、htd3和互补转基因T2代植株分蘖期表型,标尺为6 cm;F~G-野生型、htd3和互补转基因T2代植株分蘖期的分蘖数(F)和株高(G)统计,数据为平均值±标准差(n=5)。用新复极差法进行多重比较,相同小写字母表示在0.05水平上无显著差异。
Fig. 3. Map-based cloning of HTD3 and verification of complementary transgene. A, Initial location of HTD3; B, Fine mapping of HTD3; C, Predicted ORFs in the location interval; D, Gene structure of the candidate gene LOC_Os12g21710 and the sequence difference between WT and htd3, in which black boxes, white boxes, black lines represent exons, UTR and introns, respectively; E, Phenotype of WT, htd3 and complementary transgenic T2 generation plants in the tilling stage, bar=6 cm; F~G, Number of tillers (F) and plant height (G) of WT, htd3 and complementary transgenic T2 generation plants in the tillering stage, the data are means ± standard deviation (n=5). Significant difference by Duncan’s multiple range test. The same letters indicate no significant difference at P<0.05.
图4 HTD3和激素相关基因的表达水平分析 A-HTD3基因在野生型和突变体htd3各组织中相对表达水平;B~C-独脚金内酯(B)和ABA(C)相关基因在野生型和突变体htd3叶片中的相对表达水平。以相应基因在野生型中的表达水平作为参考并设为1, 数据为平均值±标准差, 3次生物学重复;采用t测验, *表示在0. 05水平上差异显著, **表示在0. 01水平上差异显著。
Fig. 4. Expression analysis of HTD3 and hormone related genes. A, HTD3 relative expression level in various tissues of WT and htd3; B-C, Relative expression of SLs (B) and ABA (C) related genes in WT and htd3. RNA was isolated from WT and htd3 leaves in B and C. Expression levels are represented as relative to the corresponding genes in WT (set as reference value of 1), and data are shown as means ± SD from three biological replicates. Asterisks indicate statistical significance between the WT and the mutant, as determined by Student’s t-test (*P < 0.05; **P < 0.01).
图5 HTD3的单倍型分析 A-HTD3的基因结构及147份水稻品种中HTD3的5'-UTR和编码区上核苷酸多态性, 黑色方块代表外显子, 白色方框代表UTR, 黑色方框之间的黑色线条代表内含子, Hap代表单倍型, 数字代表相对起始密码子ATG的碱基位置;B-每种单倍型对应的分蘖数和株高。用新复极差法进行多重比较, 相同小写字母表示在0.05水平上无显著差异。
Fig. 5. Analysis of the HTD3 gene haplotypes. A, Gene structure of the HTD3, and nucleotide polymorphisms in the HTD3 5'-UTR and coding region, exons are indicated by black boxes, UTRs are indicated by white boxes, and the black lines between black boxes represent introns, hap means haplotypes, the number represents the base position relative to the start codon ATG. B, Tiller number and plant height of each haplotype. Significant difference by Duncan’s multiple range test. The same letters indicate no significant difference at P<0.05.
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