谷子条纹叶突变体A36-S的细胞学特性分析及基因定位

doi:10.3864/j.issn.0578-1752.2021.14.003

中国农业科学 ›› 2021, Vol. 54 ›› Issue (14): 2952-2964.doi: 10.3864/j.issn.0578-1752.2021.14.003

• 作物遗传育种·种质资源·分子遗传学 • 上一篇下一篇

谷子条纹叶突变体A36-S的细胞学特性分析及基因定位

张硕^1,²(),智慧¹,唐婵娟¹,罗明昭¹,汤沙¹,贾冠清¹,贾彦超¹,刁现民¹()

¹中国农业科学院作物科学研究所,北京 100081
²湖北省农业科学院粮食作物研究所,武汉 430064

收稿日期:2020-12-11 接受日期:2021-02-01 出版日期:2021-07-16 发布日期:2021-07-26
通讯作者: 刁现民
作者简介:张硕,E-mail： zhangshuo0728@126.com。
基金资助:
国家重点研发计划(2019YFD1000704);国家重点研发计划(2019YFD1000700);中国农业科学院科技创新工程协同创新任务“作物高光效育种生物学基础及核心材料创制”(CAAS-XTCX2016002);国家现代农业产业技术体系(CARS-06-13.5-A4);中国农业科学院科技创新工程(特色农作物优异种质资源发掘与创新利用创新团队);湖北省农科院青年科学基金(2020NKYJJ02)

Cytological Characters Analysis and Low-Resolution Mapping of Stripe-Leaf MutantA36-S in Foxtail Millet

ZHANG Shuo^1,²(),ZHI Hui¹,TANG ChanJuan¹,LUO MingZhao¹,TANG Sha¹,JIA GuanQing¹,JIA YanChao¹,DIAO XianMin¹()

¹Institute of Crop Science, Chinese Academy of Agricultural Science, Beijing 100081
²Food Crop Research Institute, Hubei Academy of Agricultural Science, Wuhan 430064

Received:2020-12-11 Accepted:2021-02-01 Online:2021-07-16 Published:2021-07-26
Contact: XianMin DIAO

摘要/Abstract

摘要：

【目的】谷子是C₄模式植物,其叶色突变体是研究C₄光合途径的良好材料。通过研究谷子条纹叶突变体A36-S的细胞学特性并对突变基因进行定位,为克隆突变基因、解析谷子叶绿体合成及发育机理、进一步理解C₄光合调控机制奠定基础。【方法】谷子条纹叶突变体A36-S是由育种创制的中间材料A36自然变异而来。对比A36-S及其正常表型等基因系A36-N的表型特征,调查二者的株高、叶宽、叶长、穗重、千粒重、结实率等农艺性状指标;测定A36-S和A36-N的叶绿素含量、净光合速率、胞间CO₂浓度、气孔导度、蒸腾速率等光合指标,分析A36-S的光合特性;观察A36-S和对照品种豫谷1号的叶片半薄横截切片和超薄切片,分析A36-S叶片解剖结构特征,分别统计叶肉细胞和维管束鞘细胞中叶绿体的数量和面积,从而分析叶绿体合成及发育情况;构建A36-S×SSR41的F₂分离群体,统计群体中正常表型单株与条纹叶单株的数量,进行遗传分析;分别构建F₂分离群体正常单株与条纹叶单株的DNA混池,采用集团分离分析法（BSA法）进行突变基因的定位;筛选、开发多个SSR标记及In-Del标记,扫描F₂群体中条纹叶单株,进行进一步基因定位。【结果】谷子条纹叶突变体A36-S在全生育期表现出叶片不规则白色条纹的表型。农艺性状分析表明,相比其近等基因系A36-N,A36-S在株高、叶宽、穗重、千粒重、结实率等表型上均显著下降。光合指标测定表明A36-S叶片中叶绿素含量明显降低,尤其是叶绿素b含量下降更为严重,同时净光合速率也明显下降。叶片解剖结构观察发现,与对照豫谷1号相比,A36-S的Kranz结构变化并不明显,但叶绿体数量和大小都显著低于对照。观察叶绿体超微结构,发现A36-S的不同细胞间叶绿体发育状况差异较大,依据叶绿体发育情况可将叶片细胞可分为3类：Ⅰ类细胞具有正常发育的叶绿体;Ⅱ类细胞叶绿体基粒及片层结构减少;Ⅲ类细胞则叶绿体结构严重异常甚至不含有叶绿体。遗传分析表明A36-S表型受隐性单基因控制,利用F₂分离群体将突变基因定位在第4染色体7.66—27.90 Mb区间内。【结论】谷子A36-S条纹叶突变体表现为农艺性状及光合能力下降,叶片细胞叶绿体的数量、大小及结构均表现出显著异常。条纹叶性状受隐性单基因控制,利用分子标记将候选基因定位于第4染色体7.66—27.90 Mb区间内。

关键词: 谷子, 条纹叶突变体, 叶绿体, 基因定位

Abstract:

【Objective】 Foxtail millet is a C₄ model plant, and its leaf color mutants are good materials for C₄ photosynthetic pathway research. Through the cytological characters analysis and gene initial mapping of the stripe-leaf mutant A36-S in foxtail millet, it laid the foundation for cloning the mutant gene, analyzing the chloroplast biogenesis and development, and further understanding the C₄ photosynthetic regulation mechanism in foxtail millet. 【Method】 The stripe-leaf mutant of foxtail millet A36-S was naturally mutated from intermediate material A36 created by breeding. Comparing the phenotypic characteristics of A36-S and its isogenic line A36-N, which showed normal phenotypes, and investigating the agronomic traits, such as plant height, leaf width, leaf length, panicle weight, thousand-grain weight, and seed setting rate. To analyze the photosynthetic characters of A36-S, the chlorophyll content, net photosynthetic rate, intercellular CO₂ concentration, stomatal conductance, and transpiration rate of A36-S and A36-Nwere determined. By observing the leaf transverse section and ultrathin section of A36-S and the control variety Yugu1, the leaf anatomical structure characters were analyzed, by counting the numbers and areas of the chloroplasts in mesophyll cells and bundle sheath cells respectively, the chloroplast biogenesis and development were assessed. An F₂ segregation population of A36-S×SSR41 were created, and genetic analysis was conducted by counting the number of normal phenotype single plant and stripe-leaf single plant in the population. The DNA mixed pools of normal single plants and stripe-leaf single plants of the F₂ segregation population were constructed separately, and the method of Bulked Segregation Analysis (BSA) was used to locate the mutant gene. By screening the stripe-leaf plants in the F₂ generation using SSR and In-Del markers, the mutant gene were furtherly located.【Result】The stripe-leaf mutant of foxtail millet A36-S showed the phenotype of irregular white stripe-leaf in the whole growth period. Agronomic traits analysis showed that compared with its isogenic line A36-N, A36-S decreased significantly in plant height, leaf width, panicle weight, thousand-grain weight, and setting percentage. Photosynthetic index measurement showed that the chlorophyll contents of A36-S were also reduced significantly, especially the chlorophyll b content declined more severely, additionally, the net photosynthetic rate was also decreased significantly. Observation of the leaf anatomical structure showed that the chloroplasts number and area were significantly lower than that of the contrast Yugu1, while the changes in Kranz structure were not obvious. Furtherly, the ultrastructure of chloroplast was observed and showed that the chloroplast development situation in different cells was quite different. And the leaf cells ofA36-S could be classified into three types: type Ⅰ cells had normal chloroplasts, type Ⅱ cells had chloroplasts with reduced grana and lamellar structures, while type Ⅲ cells had severely abnormal chloroplasts or even had no chloroplast. Genetic analysis suggested that the stripe-leaf trait of A36-S was controlled by a single recessive gene, and the mutant gene was located to the region from 7.66 Mb to 27.90 Mb of chromosome 4 by F₂ population. 【Conclusion】 Stripe-leaf mutant of foxtail millet A36-S represented decreased agronomic traits and photosynthetic capacity, and the number, size, and ultrastructure of leaf cell chloroplast were significantly abnormal. The stripe-leaf trait of A36-S was controlled by a single recessive gene, which was mapped to a region from 7.66 Mb to 27.90 Mb of chromosome 4.

Key words: Setaria italica (L.) P. Beauv., stripe-leaf mutant, chloroplast, gene mapping

张硕,智慧,唐婵娟,罗明昭,汤沙,贾冠清,贾彦超,刁现民. 谷子条纹叶突变体A36-S的细胞学特性分析及基因定位[J]. 中国农业科学, 2021, 54(14): 2952-2964.

ZHANG Shuo,ZHI Hui,TANG ChanJuan,LUO MingZhao,TANG Sha,JIA GuanQing,JIA YanChao,DIAO XianMin. Cytological Characters Analysis and Low-Resolution Mapping of Stripe-Leaf MutantA36-S in Foxtail Millet[J]. Scientia Agricultura Sinica, 2021, 54(14): 2952-2964.

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农艺性状 Agronomic traits	A36-N	A36-S	P值 P-value	相比A36-N Compared to A36-N (%)
株高Plant height (cm)	94.58±3.00	80.32±6.42	0.001001	-15.08**
叶长Leaf length (cm)	28.42±0.86	28.48±1.39	0.468343	0.21
叶宽Leaf width (cm)	2.52±0.16	1.88±0.19	0.000239	-25.40**
茎节数Stem nodes number	11.00±0.71	10.40±0.55	0.086002	-5.45
穗长Panicle length (cm)	19.80±1.30	18.52±0.69	0.044184	-6.46*
穗粗Panicle diameter (cm)	2.98±0.13	2.84±0.19	0.109332	-4.70
单株穗数Panicle number per plant	2.20±0.84	2.20±0.84	0.500000	0.00
主穗重Weight of main panicle (g)	13.53±1.86	10.31±2.20	0.018266	-23.82*
码数Branch number	20.00±2.55	21.60±1.14	0.118035	8.00
千粒重Thousand-grain weight (g)	2.78±0.08	2.55±0.11	0.011091	-8.27*
结实率Seed setting rate (%)	68.54±3.60	42.46±10.05	0.000285	-38.34**

标记名称 Marker name	位置 Location (bp)	引物序列 Primer sequence (5’-3’)	交换株数量 Number of recombinant individuals
In4-3	2485441	F：GTATTCCCATGTTCGACGCGC R：GTCGGTTATTAGTTGACTTGTTGTG	167
InDel 14^[25]	4232606	F：TACTCATTGCATCCCCTTCAGCAGC R：CACTGGATAACGCATGGACTGACTA	105
In4-17	6847221	F：TGATGCCTTACTCCAGATTTCCAGC R：GATGCATACTCAATACTCTCTAGCA	26
In4-20	7661828	F：GACTGTGAGGAGCCGCGTGTAC R：GGCTCTCTCCCCTAACCGGC	1
In4-21	7873802	F：TGCTTCCCCAAATCAGTCTTTTAGT R：GGGACACCTTAGAAACAAAACCAGA	0
In4-29	23538740	F：AGAAGGACTGCTTTGATCTGGC R：GCCGCACACCATTTTGCATTACCT	0
In4-32	27896330	F：GCAATCGTCGTACACTACTCCA R：GTAGCCAAGTGGTCATTGTTTT	4
In4-33	30243701	F：GGTTGTCTTGTGTGTTCT R：GACAGGCTTCAAGATAAG	15
SiCAAS4034^[31]	30398635	F：CGGTGGGGTACATCGATCCTGCGACAT R：TTTTCCCTTGAATGGCACGCTGCCGGG	18

谷子条纹叶突变体A36-S的细胞学特性分析及基因定位

Cytological Characters Analysis and Low-Resolution Mapping of Stripe-Leaf MutantA36-S in Foxtail Millet

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单株号 Code	In4-3 (2 485 441 bp)	InDel 14^[25] (4 232 606 bp)	SiCAAS4019^[31] (9 999 146 bp)	SiCAAS4033^[31] (23 483 301 bp)	SiCAAS4034^[31] (30 398 635 bp)
3	H	S	S	S	S
7	H	H	S	S	S
9	S	S	S	S	H
11	H	S	S	S	S
12	H	S	S	S	S
14	H	S	S	S	S
18	H	S	S	S	S
22	H	S	S	S	S
27	H	H	S	S	S
29	H	S	S	S	S
30	H	H	S	S	S
31	H	H	S	S	S
32	H	S	S	S	S
36	H	H	S	S	S

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单株号 Code	In4-3 (2 485 441 bp)	InDel 14^[25] (4 232 606 bp)	SiCAAS4019^[31] (9 999 146 bp)	SiCAAS4033^[31] (23 483 301 bp)	SiCAAS4034^[31] (30 398 635 bp)
3	H	S	S	S	S
7	H	H	S	S	S
9	S	S	S	S	H
11	H	S	S	S	S
12	H	S	S	S	S
14	H	S	S	S	S
18	H	S	S	S	S
22	H	S	S	S	S
27	H	H	S	S	S
29	H	S	S	S	S
30	H	H	S	S	S
31	H	H	S	S	S
32	H	S	S	S	S
36	H	H	S	S	S

单株号 Code	In4-3 (2 485 441 bp)	InDel 14^[25] (4 232 606 bp)	SiCAAS4019^[31] (9 999 146 bp)	SiCAAS4033^[31] (23 483 301 bp)	SiCAAS4034^[31] (30 398 635 bp)
3	H	S	S	S	S
7	H	H	S	S	S
9	S	S	S	S	H
11	H	S	S	S	S
12	H	S	S	S	S
14	H	S	S	S	S
18	H	S	S	S	S
22	H	S	S	S	S
27	H	H	S	S	S
29	H	S	S	S	S
30	H	H	S	S	S
31	H	H	S	S	S
32	H	S	S	S	S
36	H	H	S	S	S