中国水稻科学 ›› 2017, Vol. 31 ›› Issue (3): 238-246.DOI: 10.16819/j.1001-7216.2017.6156 238
吕育松1,#, 谢耘丰1,2,#, 圣忠华1, 邬亚文1, 唐绍清1, 胡培松1, 魏祥进1,*()
收稿日期:
2016-11-24
修回日期:
2017-02-06
出版日期:
2017-05-10
发布日期:
2017-05-10
通讯作者:
魏祥进
基金资助:
Yusong LÜ1,#, Yunfeng XIE1,2,#, Zhonghua SHENG1, Yawen WU1, Shaoqing TANG1, Peisong HU1, Xiangjin WEI1,*()
Received:
2016-11-24
Revised:
2017-02-06
Online:
2017-05-10
Published:
2017-05-10
Contact:
Xiangjin WEI
摘要:
【目的】研究揭示潇湘矮矮秆小粒的遗传机制,为潇湘矮的育种利用提供理论基础。【方法】利用水稻育种过程中自然突变而得到的稳定遗传的矮秆小粒水稻材料潇湘矮,对其进行农艺性状考查、赤霉素(GA3)和油菜素内酯(BR)敏感性分析、遗传学分析,并利用潇湘矮与其近等基因系NIL(NIP)衍生的F2群体对控制矮秆小粒的基因xxa进行图位克隆,最终利用转基因互补试验验证候选基因。【结果】潇湘矮除表现为矮秆小粒外,其穗长变短,穗型紧凑,千粒重极显著下降。不同浓度梯度的赤霉素(GA3)和油菜素内酯(BR)处理,发现潇湘矮对GA3部分敏感,而对BR不敏感。遗传分析发现其符合孟德尔3:1分离规律。图位克隆将xxa基因定位于第5染色体InDel标记F81和F82之间约70 kb区间的物理距离内。该区间包含8个开放阅读框(ORF)。其中,第5个ORF(LOC_Os05g26890)被注释为水稻株高基因D1。序列分析发现,潇湘矮的D1基因在第5和12外显子分别有1个碱基的无义替换和3个碱基的缺失,其中第12外显子3个碱基的缺失导致1个赖氨酸的缺失。转基因互补显示D1可以恢复潇湘矮的表型。【结论】潇湘矮控制株高的途径可能与GA3代谢有关;潇湘矮矮秆小粒表型符合单隐性核基因控制的遗传规律;潇湘矮矮秆小粒性状是由D1基因突变所致。
中图分类号:
吕育松, 谢耘丰, 圣忠华, 邬亚文, 唐绍清, 胡培松, 魏祥进. 矮秆小粒水稻潇湘矮的形态学与分子遗传学分析[J]. 中国水稻科学, 2017, 31(3): 238-246.
Yusong LÜ, Yunfeng XIE, Zhonghua SHENG, Yawen WU, Shaoqing TANG, Peisong HU, Xiangjin WEI. Morphological and Molecular Genetic Analysis of a Dwarf and Small Grain Rice Variety Xiaoxiang’ai[J]. Chinese Journal OF Rice Science, 2017, 31(3): 238-246.
引物 Marker name | 前引物序列 Forward(5′-3′) | 后引物序列 Reverse(5′-3′) |
---|---|---|
RM18408 | GGCACTCTGGTTCCTCAATGG | TCTGTATCATCAACGGCAACTCC |
RM5140 | GGCACTCGTATTTCTCAACTTCTCC | GGGTGTATCAGGAGTACAGGTTGC |
RM18457 | ATCCTCCACCGCTCAAGAACACG | CGAGGCCATTCATCGAACAAAGC |
RM18402 | TTATGAGGCAGCCCGTAATGTTGC | GCAGCGGTGTCAACAGCTTCC |
RM18450 | AAGGCTCCATGGTTGGTTGC | CGATGGACAGACAGTGTGTAGTGG |
F78 | GGGACGAATTCTTTTCGATTAC | CGTGGACCAATTTTGGTAACTG |
F79 | AATTATTCCACTATGCACATGT | ATTTTCTTCCATCGCCTCTTGC |
F81 | GTAAACTATCGACTTGCTATGT | ACTAGTGCAGACTGTTTTCCTG |
F82 | AGATGATTGGATGAGAATTTAA | ACCCAGAAACCATCTAGTAATT |
F84 | GGGTGGCTCCTTACGACATTAC | TTTCATATTTTAGCGGTGCTCT |
F85 | CCTCGACCCACTGCATCATCAG | TGGTCGGTCTCCTCCCTCTTCA |
F86 | CAGTATTCGAGTAAGTTCACAA | CCTCGCCCCTATTCATCCTCTT |
表1 本研究中基因定位所用引物
Table 1 Primers for mapping in the study.
引物 Marker name | 前引物序列 Forward(5′-3′) | 后引物序列 Reverse(5′-3′) |
---|---|---|
RM18408 | GGCACTCTGGTTCCTCAATGG | TCTGTATCATCAACGGCAACTCC |
RM5140 | GGCACTCGTATTTCTCAACTTCTCC | GGGTGTATCAGGAGTACAGGTTGC |
RM18457 | ATCCTCCACCGCTCAAGAACACG | CGAGGCCATTCATCGAACAAAGC |
RM18402 | TTATGAGGCAGCCCGTAATGTTGC | GCAGCGGTGTCAACAGCTTCC |
RM18450 | AAGGCTCCATGGTTGGTTGC | CGATGGACAGACAGTGTGTAGTGG |
F78 | GGGACGAATTCTTTTCGATTAC | CGTGGACCAATTTTGGTAACTG |
F79 | AATTATTCCACTATGCACATGT | ATTTTCTTCCATCGCCTCTTGC |
F81 | GTAAACTATCGACTTGCTATGT | ACTAGTGCAGACTGTTTTCCTG |
F82 | AGATGATTGGATGAGAATTTAA | ACCCAGAAACCATCTAGTAATT |
F84 | GGGTGGCTCCTTACGACATTAC | TTTCATATTTTAGCGGTGCTCT |
F85 | CCTCGACCCACTGCATCATCAG | TGGTCGGTCTCCTCCCTCTTCA |
F86 | CAGTATTCGAGTAAGTTCACAA | CCTCGCCCCTATTCATCCTCTT |
图1 潇湘矮(XXA)及其近等基因系NIL(NIP)的表型分析 A—NIL(NIP)(左)与潇湘矮(右)5周大小苗期表型;B—成熟期NIL(NIP)(左)和潇湘矮(右)植株表型;C—成熟期NIL(NIP)(左)和潇湘矮(右)主穗表型;D—NIL(NIP)(上)和潇湘矮(下)粒型比较;E-J—NIL(NIP)与潇湘矮株高、每株穗数、结实率、千粒重、粒长以及粒宽比较。A~C图中标尺为10 cm,D图中标尺为5 mm;E~J中数据为10次重复实验的平均值,利用t测验检测数据差异显著性(*P< 0.05, **P< 0.01)。
Fig. 1. Phenotype analysis of XXA and NIL(NIP). A, 5-weeks-old seedlings of NIL(NIP)(left) and XXA(right); B, Phenotypes of NIL(NIP) and XXA plants at the mature stage in paddy field; C, Comparison of main panicles between NIL(NIP) and XXA at the mature stage; D, Grain size of NIL(NIP) and XXA at the mature stage; E-J, Plant height(E), number of panicles per plant(F), seed setting rate(G), 1,000-grain weight(H), grain length(I) and grain width(J) between NIL(NIP) and XXA. Bar=10 cm in Figures A, B and C; Bar=5 mm in Figure D. Data in E-J are shown as Mean±SD from 10 individual replicates. The asterisks indicate statistical significance, as determined by the Student’s t-test(*P<0.05, **P<0.01).
图2 潇湘矮与近等基因系NIL(NIP)主茎穗长、节间长度 A–近等基因系NIL(NIP)(左)与潇湘矮(右)主茎不同节间及穗长比较,标尺为15 cm;B–近等基因系NIL(NIP) 与潇湘矮主茎不同节间长度及穗长差异统计分析(Mean±SD, **P<0.01, n=10,t测验)。
Fig. 2. The internode and panicle length comparison in XXA and NIL(NIP). A, Differences in panicles and internode length for NIL(NIP)(left) and XXA(right) plants. P, Panicle. Those from I to V indicate the corresponding internodes from the top to bottom, bar = 15 cm; B, Internode and young panicle length analysis of NIL(NIP) and XXA at the heading stage. Data in B are shown as Mean±SD from 10 individual replicates. The asterisks indicate statistical significance, as determined by the t-test(**P<0.01).
图3 潇湘矮与NIL(NIP) 幼苗期第2叶叶鞘对外源赤霉素(A)和油菜素内酯(B)的响应 A, B中数据为10次重复实验的平均值±标准差。
Fig. 3. Elongation of the second leaf sheath in response to gibberellin(A) and brassinolide(B) in NIL(NIP) and XXA plants. Data in A, B are shown as Mean±SD from 10 individual replicates.
图4 矮秆小粒基因的精细定位与候选基因分析 A–潇湘矮矮秆小粒基因的精细定位;B–候选基因 LOC_Os05g26890的结构及其在潇湘矮中的突变位点。箭头指示位置分别为1个碱基突变和3个碱基缺失,前者为无义突变,后者造成1个赖氨酸缺失。
Fig. 4. Fine mapping and positional cloning of the candidate gene. A, Fine mapping of the xxa gene; B, The structure of candidate gene LOC_Os05g26890 and the mutation sites in XXA. Black arrows show 1-bp nonsense mutation in the 5th exon and a 3-bp deletion that resulted in a lysine deletion in the 12th exon of D1.
F1组合 Combination of F1 | 总植株数 Total No. of plants | 正常植株数 No. of half-dwarfing plants | 矮秆小粒植株数 No. of dwarfing plants | 理论比 Expressed ratio | χ2 |
---|---|---|---|---|---|
日本晴/潇湘矮 Nipponbare/XXA | 3223 | 2341 | 782 | 3:1 | 0.078 |
热研1号/潇湘矮 Reyan 1/XXA | 2964 | 2249 | 725 | 3:1 | 0.420 |
0248/潇湘矮 0248/XXA | 3744 | 2812 | 932 | 3:1 | 0.937 |
Y01/潇湘矮 Y01/XXA | 3416 | 2560 | 856 | 3:1 | 0.937 |
表2 日本晴、热研1号、0248、Y01和潇湘矮杂交组合F2株高的分离情况
Table 2 Segregation of combination Nipponbare/XXA , Reyan 1/XXA, 0248/XXA and Y01/XXA.
F1组合 Combination of F1 | 总植株数 Total No. of plants | 正常植株数 No. of half-dwarfing plants | 矮秆小粒植株数 No. of dwarfing plants | 理论比 Expressed ratio | χ2 |
---|---|---|---|---|---|
日本晴/潇湘矮 Nipponbare/XXA | 3223 | 2341 | 782 | 3:1 | 0.078 |
热研1号/潇湘矮 Reyan 1/XXA | 2964 | 2249 | 725 | 3:1 | 0.420 |
0248/潇湘矮 0248/XXA | 3744 | 2812 | 932 | 3:1 | 0.937 |
Y01/潇湘矮 Y01/XXA | 3416 | 2560 | 856 | 3:1 | 0.937 |
图5 转基因功能互补 A~D–潇湘矮(左)和转基因阳性T0家系(D1/XXA-1, 右) 成熟期表型。包括株型(A)、分枝(B)、粒型(C, 上为D1/XXA-1, 下为XXA)、节间以及穗长(D); E–D1基因在潇湘矮(XXA)以及转基因互补家系中表达水平。F~I–潇湘矮与互补转基因植株在节间长度(F)、株高(G)、粒长(H)以及粒宽(I)比较。A, B, D图中Bar = 10 cm,C图中Bar = 5 mm;E中数据为3次重复实验的平均值,F~I中数据为10次重复实验的平均值,利用t测验检测数据差异显著性(*P <0.05, **P <0.01)。
Fig. 5. Phenotype analysis of complementary plants(D1/XXA). A-D, Phenotypes of the complementary plants of D1/XXA-1(right) and XXA(left) at harvesting time, including plant type(A), panicle(B), grain size(C) and internode(D) of D1/XXA-1 and XXA; E, Relative expression levels of D1 gene in XXA and D1/XXA lines. F-I, Internode length(F), plant height(G), grain length(H) and grain width(I) of D1/XXA-1 and XXA(Bar=10 cm in A, B and D; Bar=5 mm in C). Data in E are shown as Mean±SD from 3 individual replicates. Data in F-I are shown as Mean±SD from 10 individual replicates. The asterisks indicate statistical significance, as determined by the Student’s t-test(*P<0.05; **P<0.01).
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