一个水稻低温移栽白条纹突变体wltt的鉴定和基因定位

doi:10.16819/j.1001-7216.2018.8026

中国水稻科学 ›› 2019, Vol. 33 ›› Issue (1): 1-11.DOI: 10.16819/j.1001-7216.2018.8026

• 研究论文 • 下一篇

一个水稻低温移栽白条纹突变体wltt的鉴定和基因定位

林添资^1,², 孙立亭², 龚红兵², 王益华¹, 刘玲珑¹, 赵志刚¹, 江玲¹, 万建民^1,^*()

¹南京农业大学作物遗传与种质创新国家重点实验室/农业部长江中下游粳稻生物学与遗传育种重点实验室/长江流域杂交水稻协同创新中心/江苏省现代作物生产协同创新中心, 南京 210095
²江苏丘陵地区镇江农业科学研究所, 江苏句容 212400

收稿日期:2018-03-13 修回日期:2018-05-29 出版日期:2019-01-10 发布日期:2019-01-10
通讯作者: 万建民
基金资助:
国家重点研发计划资助项目(2017YFD0100400); 江苏省重点研发计划资助项目(BE2015363); 江苏省农业科技自主创新资金资助项目[CX(16)1029]

Identification and Gene Mapping of a white-stripe leaf after transplanting at low temperature Mutant in Rice

Tianzi LIN^1,², Liting SUN², Hongbin GONG², Yihua WANG¹, Linglong LIU¹, Zhigang ZHAO¹, Ling JIANG¹, Jianmin WAN^1,^*()

¹State Key Laboratory of Crop Genetics and Germplasm Enhancement, Nanjing Agricultural University/Key Laboratory of Biology, Genetics and Breeding of japonica Rice in Mid-lower Yangtze River, Ministry of Agriculture/The Yangtze River Valley Hybrid Rice Collaboration Innovation Center/Jiangsu Collaboration Innovation Center for Modern Crop Production, Nanjing 210095, China
²Zhenjiang Agricultural Research Institute, Jurong 212400, China

Received:2018-03-13 Revised:2018-05-29 Online:2019-01-10 Published:2019-01-10
Contact: Jianmin WAN

摘要/Abstract

摘要：

【目的】叶色突变相关基因的鉴定与克隆为研究叶绿体发育、叶绿素合成和光合作用等分子机制提供理论基础。【方法】从常规粳稻镇糯19杂交后代中分离出一个低温移栽后叶色转成白条纹的自然变异突变体, 命名为wltt (white stripe leaf after transplanting at low temperature)。成熟期测定野生型和wltt的主要农艺性状,分别在苗期、移栽后15 d和同时期直播条件下测定新生叶片的色素含量并观察叶绿体的超微结构;将wltt和野生型正反交进行遗传分析;用wltt与籼稻9311杂交产生的F₂作为定位群体进行基因定位;采用RT-qPCR 分析叶绿体发育、叶绿素合成和光合作用相关基因在野生型和wltt中的表达水平。【结果】wltt突变体在苗期表现正常绿色,移栽15 d后心叶出现白条纹叶表型,至分蘖末期心叶叶色恢复;而不经移栽,突变体不会出现白条纹叶。人工模拟实验表明该表型是由低温条件下根损伤引起的。与野生型相比,wltt突变体移栽后的新生叶色素含量显著降低,光合速率下降;同时株高变矮,穗长、剑叶长和每穗粒数均显著降低。叶绿体的超微结构显示,突变体的叶肉细胞中,仅少数细胞含有正常的叶绿体,其余大部分叶肉细胞不含叶绿体。进一步研究发现,突变体中部分光合系统相关基因和叶绿体发育相关基因表达下调,叶绿素生物合成相关的14个基因表达也下调。遗传分析表明,该突变性状受一对隐性核基因控制。利用wltt突变体/9311的F₂群体,将该基因定位于水稻第2染色体着丝粒附近853 kb区间内。目前,该区间内没有叶色相关基因的报道。【结论】WLTT是低温条件下移栽调控叶片转色的关键基因,在叶绿体发育过程中发挥重要作用。

关键词: 水稻, 移栽叶片转色, 叶绿体发育, 基因定位

Abstract:

【Objective】 Isolation and characterization of leaf-color mutation related genes lays a firm theoretical foundation for dissecting the molecular mechanism underlying chloroplast development, chlorophyll biosynthesis, and photosynthesis in rice. 【Method】A spontaneous leaf-color mutant, termed as white stripe leaf after transplanting at low-temperature (wltt), was obtained from the progeny of japonica cultivar Zhennuo 19. The main agronomic traits of the wild type and wltt were determined at maturity. The pigment contents and ultrastructure of chloroplast of newly emerged leaves were analyzed at the seedling stage, fifteen days after transplanting, at the tillering stage under direct seeding. Genetic analysis was carried out by reciprocal cross of the wild type and wltt. An F₂ population derived from the cross wltt×9311 was used for gene mapping. Quantitative RT-PCR was carried out to analyze the relative expression of genes associated with chloroplast development and chlorophyll biogenesis in the wild type and the wltt mutant. 【Result】The white-striped leaves in the wltt mutant only emerged at 15 days after transplanting at low temperature such as 20℃. No white-striped leaf was observed under direct seeding treatment. However, leaves of the mutant developed normally at the late tillering stage. Simulation experiments showed that the mutant phenotype was caused by root injury at low temperature. Compared with the wild type, the pigment contents in white-stripe leaves of the wltt mutant were significantly decreased, accompanying by reduced photosynthetic rate. Simultaneously, most of the mesophyll cells had no chloroplasts. The expression levels of genes associated with chloroplast development, chlorophyll biosynthesis, and photosynthetic system were all down-regulated in the mutant. At maturity, the mutant was featured with reduced plant height, panicle length, flag leaf length and number of spikelets per panicle relative to its wild type. Genetic analysis revealed that the mutant phenotype was controlled by a single recessive nuclear gene. Moreover, the WLTT gene was mapped within an 853 kb region near the centromere on chromosome 2, between InDel markers L22 and L26, in which no gene related to leaf color was reported. 【Conclusion】WLTT is a key gene regulating leaf color after transplanting at low temperature, which plays an important role in chloroplast development.

Key words: rice, white-stripe leaf after transplanting, chloroplast development, gene mapping

中图分类号:

林添资, 孙立亭, 龚红兵, 王益华, 刘玲珑, 赵志刚, 江玲, 万建民. 一个水稻低温移栽白条纹突变体wltt的鉴定和基因定位[J]. 中国水稻科学, 2019, 33(1): 1-11.

Tianzi LIN, Liting SUN, Hongbin GONG, Yihua WANG, Linglong LIU, Zhigang ZHAO, Ling JIANG, Jianmin WAN. Identification and Gene Mapping of a white-stripe leaf after transplanting at low temperature Mutant in Rice[J]. Chinese Journal OF Rice Science, 2019, 33(1): 1-11.

图/表 11

表1 基因定位引物

Table 1 Primers used for gene mapping.

引物名称 Primer name	前引物 Forward sequence (5′-3′)	后引物 Reverse sequence (5′-3′)	BAC克隆 BAC clone
L1	ATTCAGTAAGACTACACGCAT	AATGACAGATTACTTGTTCCA	OJ1756_H07
L6	CTAACATAATGGGTAAAGAGG	TTAGTTGGTTGCCGTGT	OJ1124_E11
L8	ATAGTTTAGGGAGTTATGTGCT	CGTGTGCCTATTGACTTCTC	OSJNBa0030M21
L11	ACAGAACGGAACGGGATA	CTCACAATCTTTTATCACCCA	OSJNBa0078K05
L14	AACCAAGAATCGGAAAGAA	ATCCCATTTCCATTTCTCT	OSJNBa0008C07
L16	TTTCCTGAGCGAATCCA	AAAAGGCACTTATGAGACACT	OSJNBb0080M22
L18	TAGGTGGTTGAATGGTGC	TATGCTTCTTTTGGGTTG	P0543C11
L20	TGAGATACGCAGAATGGG	GAGGAGGATGCAGGGAC	P0705A04
L22	GTTCTTTTGTCTTCCCTCA	ATTATCCTTGGTCTTGGTAT	OJ1134_B09
L26	TTGGAGAATGAAGTTGCTAA	TTACCAAGCAGGACTAAAGAT	OSJNBb0037J12
I2-7	GAACCAGTCCGCTCTCTGAC	TACGCGTCGTGTATCGTAGC	OSJNBa0035A24

图1 野生型与wltt突变体不同时期的表型A-苗期;B-分蘖期,移栽后15 d,右上角为野生型(左)和突变体(右)的新完全展开叶;C-抽穗期移栽后15 d,右上角为野生型(左)和突变体(右)的再生分蘖的叶片;D-6月10日正常移栽的成熟期植株;E-直播条件下分蘖期的植株;F-正常移栽分蘖期植株新生叶片的净光合速率。WT-野生型。**表示经t测验后,P<0.01下差异显著。

Fig. 1. Phenotypic characterization of the wltt mutant and its wild type(WT). A, Seedling stage; B, Fifteen days after transplanting, at the tillering stage; The insert represents the new fully-expanded leaf blades of the wild type (left) and the mutant (right); C, Fifteen days after transplanting at the heading stage. The inserts represents the leaves of regenerated tillers of the wild type (left) and the mutant (right); D, Plants at the mature stage (transplanting at the tillering stage). E, Wild-type and mutant plants at the tillering stage under direct seeding. F, Net photosynthetic rate of new fully expanded leaf blades of the wild type and wltt mutant at the tillering stage. WT, Wild type. ** P<0.01(Student’s t test).

表2 野生型与突变体wltt的主要农艺性状比较

Table 2 Comparison of major agronomic traits between the wltt mutant and its wild type(WT).

材料 Material	株高 Plant height /cm	有效穗数 No. of effective panicles	剑叶长 Flag leaf length /cm	穗长 Panicle length /cm		每穗总粒数 No. of spikelets per panicle	结实率 Seed-setting rate /%	千粒重 1000-grain weight /g
WT	86.7±0.2	9.2±0.4	22.28±0.53		20.44±0.46	173.6±5.7	95.8±0.4	29.18±0.26
wltt	75.8±0.5^**	8.2±0.4	18.10±0.37^**		18.40±0.41^**	136.2±6.9^**	95.6±1.7	28.46±0.32

图2 wltt突变体和野生型叶片色素含量分析 Chla-叶绿素a;Chlb-叶绿素b;Car-类胡萝卜素;Total-总色素含量。误差线表示3次独立实验的标准差。**表示野生型与突变体间差异达0.01显著水平（t测验）。

Fig. 2. Pigment contents in leaves of wltt mutant and its wild type(WT). Chla, Chlorophyll a; Chlb, Chlorophyll b; Car, Carotenoids; Total, Total pigment contents. Values are presented as mean±SD. **The difference between the wild type and wltt is significant at 0.01 level according to Student’s t test.

图3 野生型和突变体wltt的叶绿体超微结构 A~C为野生型的叶绿体超微结构;D~F为突变体wltt中同时含有异常结构的叶绿体(D)和正常结构的叶绿体(E~F)。Cp-叶绿体; Thy-类囊体; OB-嗜锇体。

Fig. 3. Transmission electron microscopic (TEM) images of chloroplast ultrastructure in the wild type and the wltt mutant. A-C, Transmission electron microscopic (TEM) images of chloroplast ultrastructure in wild type; D-F, Two types of chloroplasts with (E, F) or without (D) a normal ultrastructure in the wltt mutant; Cp, Chloroplast; Thy, Thylakoid lamellae; OB, Osmiophilic body.

图4 叶绿体发育及光合系统相关基因在野生型和wltt突变体中的表达分析误差线表示3次独立实验的标准差。**表示野生型与突变体间差异达0.05和0.01显著水平（t测验）。

Fig. 4. Expression analysis of genes associated with chloroplast development and photosynthetic system in the wild type(WT) and wltt mutant. Mean±SD (n=3). **The difference between the wild type and wltt is significant at 0.01 level according to Student’s t test.

图5 叶绿素合成相关基因在野生型和突变体wltt中的表达分析误差线表示3次独立实验的标准差。*和**分别表示野生型与突变体间差异达0.05和0.01显著水平（t测验）。

Fig. 5. Expression analysis of genes associated with chlorophyll biogenesis in the wild type(WT) and wltt mutant. Means ± SD (n=3). * and **The difference between the wild type and wltt is significant at 0.05 and 0.01 level, respectively according to Student’s t test.

图6 环境条件对叶色变异的影响 A–不进行剪根处理;B, C-剪根处理10 d后,20℃条件下,光照强度为25 000 lx(B)和6250 lx(C);D, E-剪根处理10 d后,30℃条件下,光照强度为25 000 lx(D)和6250 lx(E)。

Fig. 6. Effects of environmental conditions on leaf color variation. A, The seedlings of the wild type and wltt with non-injured roots; B, C, The seedlings of wild type and wltt 10 days after root cutting treatment at 20℃ at the light intensity of 25 000 lx(B) and 6250 lx(C), respectively. D, E, The seedlings of wild type and wltt 10 days after root cutting treatment at 30℃ at the light intensity of 25 000 lx(D) and 6250 lx(E), respectively.

图7 环境条件对剪根处理后色素含量的影响误差线表示3次独立实验的标准差。**表示野生型与突变体间差异达0.01显著水平（t测验）。

Fig. 7. Effects of environmental conditions on pigment contents after root cutting treatment. Means±SD (n=3). **The difference between the wild type and wltt is significant at 0.01 level according to Student’s t test.

表3 野生型与wltt突变体的F2代分离比

Table 3 Segregation of F2 population from wltt mutant and its wild type(WT).

杂交组合 Cross	正常株数 No. of normal plants	白条纹株数 No. of white stripe plants	实际分离比 Segregation ratio	χ²_(3:1)
WT×wltt	375	127	2.95:1	0.02
wltt×WT	368	130	2.83:1	0.06

图8 WLTT在第2染色体上的定位 A–基因位点初定位在第2染色体着丝粒附近InDel标记I2-5和I2-8之间;B-基因位点精细定位到标记L22和L26之间,物理距离853 kb。n为定位所用的白条纹叶植株数。

Fig. 8. Location of WLTT on rice chromosome 2. A, The gene is mapped to the centromeric region of chromosome 2 between the InDel markers I2-5 and I2-8; B, Mapping of the gene locus between markers L22 and L26 within an 853 kb region. CEN, Centromere; n, Number of individuals with white stripe leaf after transplanting.

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一个水稻低温移栽白条纹突变体wltt的鉴定和基因定位

Identification and Gene Mapping of a white-stripe leaf after transplanting at low temperature Mutant in Rice

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