生物技术通报 ›› 2024, Vol. 40 ›› Issue (7): 125-136.doi: 10.13560/j.cnki.biotech.bull.1985.2024-0132
庞梦真1,2(), 徐汉琴1,2, 刘海燕1,2, 宋娟1,2, 王佳涵1,2, 孙丽娜1,2, 姬佩梅1,2, 尹泽芝1,2, 胡又川1,2, 赵晓萌1,2, 梁闪闪1,2, 张泗举1,2(), 栾维江1,2()
收稿日期:
2024-02-02
出版日期:
2024-07-26
发布日期:
2024-05-24
通讯作者:
栾维江,男,博士,教授,研究方向:水稻功能基因组学;E-mail: lwjzsq@163.com;作者简介:
庞梦真,女,硕士研究生,研究方向:遗传学;E-mail: mengzhenpang@163.com
基金资助:
PANG Meng-zhen1,2(), XU Han-qin1,2, LIU Hai-yan1,2, SONG Juan1,2, WANG Jia-han1,2, SUN Li-na1,2, JI Pei-mei1,2, YIN Ze-zhi1,2, HU You-chuan1,2, ZHAO Xiao-meng1,2, LIANG Shan-shan1,2, ZHANG Si-ju1,2(), LUAN Wei-jiang1,2()
Received:
2024-02-02
Published:
2024-07-26
Online:
2024-05-24
摘要:
【目的】 水稻的抽穗期对水稻地域适应性及水稻产量至关重要,对水稻抽穗期基因进行鉴定及功能分析,可以为水稻育种提供优异的基因资源。【方法】 通过BSA-seq方法对一个黄化早抽穗突变体hz1(huangzao 1)进行基因定位克隆及连锁分析;利用RT-qPCR技术分析目的基因HZ1的表达谱,并用水稻原生质体瞬时转化查明HZ1的亚细胞定位;对突变体的抽穗期、叶绿素含量、过氧化氢含量等生理指标进行测定,详细分析其表型变化。【结果】 田间表型观察发现hz1表现早抽穗,长日照(long-day, LD)及短日照(short-day, SD)条件下hz1的抽穗期相同,分别比野生型(wild type, WT)早抽穗43 d和26 d,表明hz1是一个光周期不敏感的突变体。同时hz1表现黄化表型,相比WT,叶绿素含量下降。遗传分析表明hz1由隐性单基因控制,F2混池高通量测序将目的基因定位于水稻第6染色体上17.8 Mb区间内,分析发现该区间内一个T-DNA插入位点LOC_Os06g40080与hz1目标性状完全连锁,LOC_Os06g40080为已知的SE5基因,编码血红素加氧酶1(heme oxygenase 1, HO1)。HZ1/SE5在叶片中高表达,在LD及SD条件下具有昼夜节律性表达。亚细胞定位发现HZ1/SE5蛋白定位于叶绿体中。表达调控分析表明HZ1/SE5主要通过调控水稻成花素Hd3a和RFT1的表达来调控水稻的抽穗期;并通过调控叶绿素合成途径相关基因的表达水平影响水稻叶绿素水平变化。【结论】 黄化早抽穗突变体hz1由于血红素加氧酶编码基因SE5突变导致其对光周期不敏感,HZ1/SE5基因通过调控水稻成花素基因及叶绿素合成途径相关基因的表达而影响水稻的抽穗期及叶片的黄化。
庞梦真, 徐汉琴, 刘海燕, 宋娟, 王佳涵, 孙丽娜, 姬佩梅, 尹泽芝, 胡又川, 赵晓萌, 梁闪闪, 张泗举, 栾维江. 水稻黄化早抽穗突变体 hz1 的基因鉴定及功能分析[J]. 生物技术通报, 2024, 40(7): 125-136.
PANG Meng-zhen, XU Han-qin, LIU Hai-yan, SONG Juan, WANG Jia-han, SUN Li-na, JI Pei-mei, YIN Ze-zhi, HU You-chuan, ZHAO Xiao-meng, LIANG Shan-shan, ZHANG Si-ju, LUAN Wei-jiang. Gene Identification and Functional Analysis of Yellowish and Early Heading Mutant hz1 in Rice[J]. Biotechnology Bulletin, 2024, 40(7): 125-136.
引物名称 Primer name | 引物序列 Primer sequence(5'-3') | 内切酶 Endonuclease |
---|---|---|
SE5-GFP-F | GTTGGTACCCGCTATAAGAGGGAGAGAGG | Kpn Ⅰ |
SE5-GFP-R | CCGTCTAGAGGTGAATATGTGACGGAGGA | Xba Ⅰ |
LB-R | ACGATGGACTCCAGTCCGGCCcttgaccaactctatcagagcttgg | |
HZ1-F | CAAGGACCAGGCCAAGGAAG | |
HZ1-R | ACCAAACTCAAAACAGGGGG | |
InDel1-F | CAGGTATTCTGAGGTTGTATCC | |
InDel1-R | CACGGTCAAATTCAACATTCC | |
dCAP3-F | CACCAACTGAGCTCACTAGCCATAT | NdeⅠ |
dCAP3-R | GGCCCAATGACTACCTTCTACTTTA | |
SE5-ex-F | GGCTGAAAAGGACTCCCAAG | |
SE5-ex-R | GTCCAGCTAGAGGCGACTTC | |
Actin1-ex-F | GACTCTGGTGATGGTGTCAGC | |
Actin1-ex-R | GGCTGGAAGAGGACCTCAGG | |
Hd3a-ex-F | TTGGTAGGGTTGTGGGTGATGTGC | |
Hd3a-ex-R | AGGTTAGGGTCACTTGGGCTTGGT | |
RFT1-ex-F | TCCGAGCCCAAGCAACCCTAAC | |
RFT1-ex-R | AGTTCCTGGTGCTGAAGTTCTG | |
HEMA-ex-F | GATGCAATCACTGCTGGAAAGCGT | |
HEMA-ex-R | CCATCTTGCCAGCACCAATCAACA | |
PORA-ex-F | TCGTCGGCCTCGTCTGAGTTTATT | |
PORA-ex-R | AGGCCTCTCTCACTGAAAGCTGAA | |
YGL1-ex-F | CCAGCCACTGATGAAAGCAGCAAT | |
YGL1-ex-R | AGAGCGCTAATACACTCGCGAACA | |
CAO1-ex-F | CTTGTCGTATTCTTGGCGAG | |
CAO1-ex-R | ATCCCGTGATGCTGTCGCTA |
表1 引物序列
Table 1 Primer sequence
引物名称 Primer name | 引物序列 Primer sequence(5'-3') | 内切酶 Endonuclease |
---|---|---|
SE5-GFP-F | GTTGGTACCCGCTATAAGAGGGAGAGAGG | Kpn Ⅰ |
SE5-GFP-R | CCGTCTAGAGGTGAATATGTGACGGAGGA | Xba Ⅰ |
LB-R | ACGATGGACTCCAGTCCGGCCcttgaccaactctatcagagcttgg | |
HZ1-F | CAAGGACCAGGCCAAGGAAG | |
HZ1-R | ACCAAACTCAAAACAGGGGG | |
InDel1-F | CAGGTATTCTGAGGTTGTATCC | |
InDel1-R | CACGGTCAAATTCAACATTCC | |
dCAP3-F | CACCAACTGAGCTCACTAGCCATAT | NdeⅠ |
dCAP3-R | GGCCCAATGACTACCTTCTACTTTA | |
SE5-ex-F | GGCTGAAAAGGACTCCCAAG | |
SE5-ex-R | GTCCAGCTAGAGGCGACTTC | |
Actin1-ex-F | GACTCTGGTGATGGTGTCAGC | |
Actin1-ex-R | GGCTGGAAGAGGACCTCAGG | |
Hd3a-ex-F | TTGGTAGGGTTGTGGGTGATGTGC | |
Hd3a-ex-R | AGGTTAGGGTCACTTGGGCTTGGT | |
RFT1-ex-F | TCCGAGCCCAAGCAACCCTAAC | |
RFT1-ex-R | AGTTCCTGGTGCTGAAGTTCTG | |
HEMA-ex-F | GATGCAATCACTGCTGGAAAGCGT | |
HEMA-ex-R | CCATCTTGCCAGCACCAATCAACA | |
PORA-ex-F | TCGTCGGCCTCGTCTGAGTTTATT | |
PORA-ex-R | AGGCCTCTCTCACTGAAAGCTGAA | |
YGL1-ex-F | CCAGCCACTGATGAAAGCAGCAAT | |
YGL1-ex-R | AGAGCGCTAATACACTCGCGAACA | |
CAO1-ex-F | CTTGTCGTATTCTTGGCGAG | |
CAO1-ex-R | ATCCCGTGATGCTGTCGCTA |
图1 hz1突变体的表型分析 A:hz1突变体在田间种植条件下的表型,比例尺为10 cm;B-D:分别为田间自然长日照(NLD)、长日照(LD)、短日照(SD)条件下WT与hz1的抽穗期,数值表示为平均值±SD(n≥25);E:苗期hz1的表型,比例尺为1 cm;F:抽穗期hz1叶片表型;G:苗期hz1的叶绿素含量测定,数值表示为平均值±SD(n=3);H:抽穗期hz1的剑叶叶绿素含量测定,数值表示为平均值±SD(n=3);I:抽穗期hz1的倒二叶叶绿素含量测定,数值表示为平均值±SD(n=3)(Chl a:叶绿素a;Chl b:叶绿素b;Chl T:总叶绿素含量);J:hz1的DAB染色结果,红色箭头所示红褐色斑点,比例尺为1 cm;K:hz1的DAB染色相对强弱分析,数值表示为平均值±SD(n=3);L:hz1突变体叶片过氧化氢含量的测定,数值表示为平均值±SD(n=3);M和N:hz1突变体叶片过氧化氢酶CAT及过氧化物酶POD的酶活性测定,数值表示为平均值±SD(n=3)。*,**,***分别表示在0.05、0.01和0.001水平上显著性差异,下同
Fig. 1 Phenotype analysis of hz1 mutant A: The phenotype of hz1 mutant in the field, bar=10 cm. B-D: The heading dates of hz1 under natural long day(NLD), long day(LD), and short day(SD)conditions. Values are shown as mean ± SD(n ≥ 25). E: The phenotype of hz1 during the seedling stage, bar=1 cm. F: The leaves of hz1 at heading stage. G: Measurement of chlorophyll content during seedling stage. Values are shown as mean ± SD(n=3). H: The chlorophyll content of flag leaves during the heading stage. Values are shown as mean ± SD(n=3). I: The chlorophyll content of the penultimate leaves. Values are shown as mean ± SD(n=3)(Chl a: chlorophyll a; Chl b: chlorophyll b; Chl T: total chlorophyll content). J: The DAB staining results of WT and hz1. Red arrows indicate brown spots, bar=1 cm. K: Relative intensity of DAB staining for WT and hz1. Values are shown as mean ± SD(n=3). L: The measurement of hydrogen peroxide(H2O2)content in leaves. Values are shown as mean ± SD(n=3). M and N: Enzyme activity measurements of catalase(CAT)and peroxidase(POD)in leaves. Values are shown as mean ± SD(n=3). *: P<0.05; **: P<0.01; ***: P<0.001, the same below
图2 hz1突变体的农艺性状分析 A-G:分别为株高、分蘖数、穗长、一次枝梗数、二次枝梗数、结实率及百粒重的统计分析,数值表示为平均值±SD(n=30);H:hz1的穗型,比例尺为10 cm;I:hz1的籽粒数及结实率,比例尺为10 cm;J和 K:hz1的花粉染色分析,放大倍数为400倍;箭头所示为典型败育花粉
Fig. 2 Analysis of agronomic traits of hz1 mutant A-G: Statistical analysis of plant height, tiller number, panicle length, number of primary branches, number of secondary branches, seed setting rate, and 100-grain weight. Values are shown as mean ± SD(n=30). H: The panicle of hz1, bar=10 cm. I: Grain number and seed setting rate of hz1, bar=10 cm. J and K: The pollen staining of hz1. The magnified times are 400 times, and the arrows indicate abortive pollens
组合 Cross | 正常表型植株数 Number of wild plants | 突变表型植株数 Number of mutant plants | 总植株数 Number of total plants | ꭕ2(3∶1) | P value |
---|---|---|---|---|---|
ZH11/hz1 | 458 | 130 | 588 | 2.621 3 | 0.105 4 |
表2 突变体hz1的遗传分析
Table 2 Genetic analysis of hz1 Mutant
组合 Cross | 正常表型植株数 Number of wild plants | 突变表型植株数 Number of mutant plants | 总植株数 Number of total plants | ꭕ2(3∶1) | P value |
---|---|---|---|---|---|
ZH11/hz1 | 458 | 130 | 588 | 2.621 3 | 0.105 4 |
图3 BSA-seq分析及基因定位 A:BSA-seq分析,HZ1与水稻第6染色体上的分子标记紧密连锁;B:T-DNA插入位点与HZ1共分离;C:T-DNA插入位点测序结果
Fig. 3 BSA-seq analysis and gene mapping A: BSA-seq analysis. HZ1 is closely linked with the molecular marker on rice chromosome 6. B: T-DNA insertion site is co-segregated with HZ1. C: Sequencing analysis of T-DNA insertion site
图4 T-DNA插入位点的检测及连锁分析 A:LOC_Os06g40080位点及T-DNA插入的位置示意图,HZ1-F、HZ1-R及LB-R所示为检测引物位置;B:HZ1/SE5基因RT-PCR表达分析,Actin1为水稻内参基因;C:T-DNA插入位点的检测及连锁分析,LD为DL 2000 marker,WT为野生型,F1为F1植株,F2 mutant和F2 WT分别为F2代群体中突变表型植株及野生型植株;D:水稻第6染色体及InDel1、dCAP3分子标记位置示意图;E和F:InDel1及dCAP3标记PCR扩增结果,LD为DL 2000 marker,P1为亲本ZH11植株,P2为亲本hz1植株,F1为F1植株,F2 mutant为F2代群体中突变表型植株
Fig. 4 Detection and linkage analysis of T-DNA insertion sites A: Schematic diagram of LOC_Os06g40080 site and T-DNA insertion location. HZ1-F, HZ1-R, and LB-R indicate the positions of the detection primers. B: RT-PCR analysis of HZ1/SE5 gene. Actin1 is an internal control in rice. C: The linkage analysis of T-DNA insertion sites. LD is DL 2000 marker, WT is wild-type, F1 is F1 plant, F2 mutant and F2 WT are plants with mutant phenotype and WT phenotype in the F2 population, respectively. D: Schematic diagram of InDel1, dCAP3 markers in the chromosome 6. E and F: PCR amplification of InDel1 and dCAP3. LD is DL 2000 marker, P1 is ZH11 plant, P2 is hz1 mutant, F1 is the F1 plants, and F2 mutant are the plants with hz1 phenotype in the F2 population
图5 HZ1/SE5基因表达模式分析 A:HZ1/SE5组织特异性表达分析(SAM:茎顶端分生组织);B:HZ1/SE5在水稻不同生长时期的表达分析;C和D分别为HZ1/SE5在LD(C)及SD(D)条件下的表达分析,ZT(zeitgeber time, h)表示给光时间。白色空心方框表示光期,黑色实心方框表示暗期。下同
Fig. 5 Expression patterns of HZ1/SE5 gene A: Tissue-specific expression analysis of HZ1/SE5(SAM: stem apical meristem). B: Expression analysis of HZ1/SE5 at different growth stages in rice. C and D: Expression analysis of HZ1/SE5 under LD (C) and SD (D) conditions, respectively. ZT(zeitgeber time, h)indicates the time of light on. The white hollow box indicates the light period, and the black solid box indicates the dark period. The same below
图6 HZ1/SE5蛋白的亚细胞定位 A和B:空载体和HZ1-GFP融合蛋白在水稻原生质体中的瞬时表达,比例尺为10 μm;C和D:空载体和HZ1-GFP融合蛋白在烟草表皮细胞中的瞬时表达,比例尺为20 μm
Fig. 6 Subcellular localization of HZ1/SE5 protein A and B: Transient expression of empty vector and HZ1-GFP fusion protein in rice protoplasts, bar=10 μm. C and D: Transient expression of empty vector and HZ1-GFP fusion protein in tobacco epidermal cells, bar=20 μm
图7 光周期调控通路关键基因与叶绿素合成相关基因在hz1突变体中的表达分析 A, B:LD及SD条件下,光周期调控通路中关键基因在WT及hz1突变体中的表达水平;C:叶绿素合成相关基因在WT与hz1突变体中的表达
Fig. 7 Expression analysis of key genes involved in photoperiodic regulatory pathway and chlorophyll synthesis related genes in hz1 mutant A, B: Expression of key genes associated with photoperiodic regulatory pathway in WT and hz1 mutants under LD and SD conditions. C: Expression of chlorophyll synthesis related genes in WT and hz1 mutants
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