小麦卷叶突变体RL1的生理特性及遗传研究

doi:10.3724/SP.J.1006.2023.31004

作物学报 ›› 2023, Vol. 49 ›› Issue (12): 3399-3410.doi: 10.3724/SP.J.1006.2023.31004

• 研究简报 • 上一篇

小麦卷叶突变体RL1的生理特性及遗传研究

刘叶¹^,²(), 李越¹, 苑名杨¹, 卫乃翠¹, 关攀锋³, 赵佳佳¹, 武棒棒¹, 郑兴卫¹, 郝宇琼¹, 乔玲¹^,^*(), 郑军¹^,^*()

¹山西农业大学小麦研究所 / 农业农村部有机旱作农业重点实验室(部省共建), 山西临汾 041000
²山西大学生命科学学院, 山西太原 030006
³郑州大学农学院, 河南郑州 450001

收稿日期:2023-01-09 接受日期:2023-06-29 出版日期:2023-12-12 网络出版日期:2023-08-04
通讯作者: * 乔玲, E-mail: qiaolingsmile@163.com; 郑军, E-mail: sxnkyzj@126.com
作者简介:刘叶, E-mail: liuye12345202@126.com^**同等贡献
基金资助:
山西农业大学省部共建有机旱作农业国家重点实验室自主研发项目(202002-1);山西省科技重大专项计划“揭榜挂帅”项目(202201140601025-2-01)

Physiological characteristics and genetic research of rolled leaf mutant1 (RL1) in wheat (Triticum aestivum L.)

LIU Ye¹^,²(), LI Yue¹, YUAN Ming-Yang¹, WEI Nai-Cui¹, GUAN Pan-Feng³, ZHAO Jia-Jia¹, WU Bang-Bang¹, ZHENG Xing-Wei¹, HAO Yu-Qiong¹, QIAO Ling¹^,^*(), ZHENG Jun¹^,^*()

¹Institute of Wheat Research, Shanxi Agriculture University / Key Laboratory of Sustainable Dryland Agriculture (Co-construction by Ministry and Province), Ministry of Agriculture and Rural Affairs, Linfen 041000, Shanxi, China
²School of Life Science, Shanxi University, Taiyuan 030006, Shanxi, China
³School of Agricultural Sciences, Zhengzhou University, Zhengzhou 450001, Henan, China

Received:2023-01-09 Accepted:2023-06-29 Published:2023-12-12 Published online:2023-08-04
Contact: * E-mail: qiaolingsmile@163.com; E-mail: sxnkyzj@126.com
About author:^**Contributed equally to this work
Supported by:
Research Program Sponsored by State Key Laboratory of Integrative Sustainable Dryland Agriculture, Shanxi Agricultural University(202002-1);Science and Technology Major Special Plan Project “Reveal the Title” of Shanxi Province(202201140601025-2-01)

摘要/Abstract

摘要：

小麦叶片在逆境下会发生可逆的折叠或卷曲, 通过脱水回避的形态学变化降低非生物胁迫的损害。目前小麦叶片卷曲的生理和遗传调控机制尚不清楚。本文利用EMS诱变晋麦47获得了卷叶突变体RL1 (Rolled Leaf 1), RL1在整个生育期叶片呈现卷曲, 初生叶片沿中轴脉向近轴面微卷, 随着叶片生长加速卷曲, 直至为筒状。与野生型相比, RL1株高降低、穗长变短、旗叶变窄和千粒重降低。氯化三苯基四氮唑(TTC)染色结果表明RL1种子活力低, 且种子发芽率降低了22%。抽穗10 d后, RL1的叶绿素含量与野生型基本一致, 净光合速率、蒸腾速率、气孔导度、细胞间隙CO₂浓度差异不显著, 但水分利用率降低。低温、高温和干旱促进RL1的叶片卷曲; 石蜡切片观察表明, RL1的大叶脉和小叶脉偏少, 在中脉区域远轴面厚壁细胞和近轴面薄壁细胞数目减少, 且维管束间泡状细胞的面积和数量均明显低于野生型; RL1叶片不同部位泡状细胞缩小以及维管束减少导致整个叶片向近轴面极度卷曲。遗传分析表明该性状受1对不完全显性的核基因控制, 位于1D染色体短臂上, 精细定位将目标区间锁定在9.42 Mb范围内。

关键词: 小麦, 卷叶突变体, 细胞学分析, 生理特性, 遗传分析

Abstract:

Wheat leaves tend to fold or curl when exposure to stresses, the dehydration avoidance in morphology can reduce the damage of abiotic stress. At present, the physiological and genetic regulation mechanism associated with leaf curling is not clear in wheat. This study reported a novel rolled leaf mutant (RL1) from the ethyl methyl sulphonate (EMS) mutagenesis cultivar Jinmai 47. The leaves of the mutant RL1 were curled during the growth period, and the primary leaves were slightly curled along axial vein to paraxial plane. Leaf curling was accelerated with the growth until the leaf was tubular. Compared to wild type (WT), plant height, ear length, flag leaf narrowing, and 1000-grain weight were decreased in mutant RL1. Triphenyltetrazolium Chloride (TTC) staining showed that seed vigor of RL1 was low, together with the decreased germination rate by 22%. Additionally, there was no significant differences in chlorophyll content, net photosynthetic rate, transpiration rate, stomatal conductance, and intercellular CO₂ concentration between RL1 and WT, while water utilization rate was decreased in RL1 after heading for 10 days. Low temperature, high temperature, and drought led to the leaf rolling in RL1. RL1 showed fewer leaf/lobular veins via paraffin section assay, and the number of abaxial sclerenchyma and adaxial parenchyma cells were reduced in midrib region of RL1. Moreover, the area and counts of vesicular cells between the vascular bundles were significantly reduced in RL1, together with the vesicular cells at the midvein region of leaves compared with WT. Vesicular cells and vascular bundles shrunk and decreased, respectively, resulting in the situation that the entire blade was extremely crimped to the adaxial plane. Genetic analysis demonstrated that the mutant trait was localized on the short arm of chromosome 1D, regulated by a pair of nuclear genes with incomplete dominance and fine mapping analysis further locked the target interval at 9.42 Mb.

Key words: wheat, rolled leaf mutant, cytological analysis, physiological characteristics, genetic analysis

刘叶, 李越, 苑名杨, 卫乃翠, 关攀锋, 赵佳佳, 武棒棒, 郑兴卫, 郝宇琼, 乔玲, 郑军. 小麦卷叶突变体RL1的生理特性及遗传研究[J]. 作物学报, 2023, 49(12): 3399-3410.

LIU Ye, LI Yue, YUAN Ming-Yang, WEI Nai-Cui, GUAN Pan-Feng, ZHAO Jia-Jia, WU Bang-Bang, ZHENG Xing-Wei, HAO Yu-Qiong, QIAO Ling, ZHENG Jun. Physiological characteristics and genetic research of rolled leaf mutant1 (RL1) in wheat (Triticum aestivum L.)[J]. Acta Agronomica Sinica, 2023, 49(12): 3399-3410.

图/表 12

表1

附表1

42对SSR引物信息(国标: NY/T 2859-2015)"

引物名称 Primer name	染色体 Chr.	退火温度 Annealing temperature (℃)	引物序列 Primer sequence (5'-3')
cwm65	1A	65	F: TCATTGGTGTCATCCCTCGTGT; R: GAATAATGCCTTGACCCTGGAC
barc80	1BL	65	F: GCGAATTAGCATCTGCATCTGTTTGAG; R: CGGTCAACCAACTACTGCACAAC
cfd72	1DL	60	F: CTCCTTGGAATCTCACCGAA; R: TCCTTGGGAATATGCCTCCT
gwm294	2AL	55	F: GGATTGGAGTTAAGAGAGAACCG; R: GCAGAGTGATCAATGCCAGA
gwm429	2BS	55	F: TTGTACATTAAGTTCCCATTA; R: TTTAAGGACCTACATGACAC
gwm261	2DS	55	F: CTCCCTGTACGCCTAAGGC; R: CTCGCGCTACTAGCCATTG
gwm155	3AL	55	F: CAATCATTTCCCCCTCCC; R: AATCATTGGAAATCCATATGCC
gwm285	3BS	65	F: ATGACCCTTCTGCCAAACAC; R: ATCGACCGGGATCTAGCC
gdm72	3DS	55	F: TGGTTTTCTCGAGCATTCAA; R: TGCAACGATGAAGACCAGAA
gwm610	4AS	65	F: CTGCCTTCTCCATGGTTTGT; R: AATGGCCAAAGGTTATGAAGG
ksum62	4B	60	F: GGAGAGGATAGGCACAGGAC; R: GAGAGCAGAGGGAGCTATGG
barc91	4DL	55	F: TTCCCATAACGCCGATAGTA; R: GCGTTTAATATTAGCTTCAAGATCAT
gwm304	5AS	55	F: AGGAAACAGAAATATCGCGG; R: AGGACTGTGGGGAATGAATG
gwm67	5BL	60	F: ACCACACAAACAAGGTAAGCG; R: CAACCCTCTTAATTTTGTTGGG
cfd29	5DL	65	F: GGTTGTCAGGCAGGATATTTG; R: TATTGATAGATCAGGGCGCA
gwm459	6AS	55	F: AATTTCAAAAAGGAGAGAGA; R: AACATGTGTTTTTAGCTATC
barc198	6BS	55	F: CGCTGAAAAGAAGTGCCGCATTATGA; R: CGCTGCCTTTTCTGGATTGCTTGTCA
cfd76	6DL	65	F: GCAATTTCACACGCGACTTA; R: CGCTCGACAACATGACACTT
cfa2028	7AS	55	F: TGGGTATGAAAGGCTGAAGG; R: ATCGCGACTATTCAACGCTT
gwm333	7BS	60	F: GCCCGGTCATGTAAAACG; R: TTTCAGTTTGCGTTAAGCTTTG
gwmn437	7DL	55	F: GATCAAGACTTTTGTATCTCTC; R: GATGTCCAACAGTTAGCTTA
wmc312	1AS	55	F: TGTGCCCGCTGGTGCGAAG; R: CCGACGCAGGTGAGCGAAG
barc240	1BL	55	F: AGAGGACGCTGAGAACTTTAGAGAA; R: GCGATCTTTGTAATGCATGGTGAAC
gdm111	1DL	55	F: CACTCACCCCAAACCAAAGT; R: GATGCAATCGGGTCGTTAGT
wmc522	2AS	55	F: AAAAATCTCACGAGTOGGGC; R: CCCGAGCAGGAGCTACAAAT
cfd51	2DS	55	F: GGAGGCTTCTCTATGGGAGG; R: TGCATCTTATCCTGTGCAGC
barc324	3AS	55	F: CCAATTCTGCCCATAGGTGA; R: GAGGAAATAAGATTCAGCCAACTG
barc164	3BS	55	F: TGCAAACTAATCACCAGCGTAA; R: CGCTTTCTAAAACTGTTCGGGATTTCTAA
cfd9	3DL	55	F: TTGCACGCACCTAAACTCTG; R: CAAGTGTGAGCGTCGG
gwm161	3DS	55	F: GATCGAGTGATGGCAGATGG; R: TGTGAATTACTTGGACGTGG
barc170	4AL	55	F: CGCTTGACTTTGAATGGCTGAACA; R: CGCCCACTTTTTACCTAATCCTTTTGAA
gwm495	4BL	55	F: GAGAGCCTCGCGAAATATAGG; R: TGCTTCTGGTGTTCCTTCG
winc720	4DS	55	F: CACCATGGTTGGCAAGAGA; R: CTGGTGATACTGCCGTGACA
gwm186	5AL	55	F: GCAGAGCCTGGTTCAAAAAG; R: CGCCTCTAGCGAGAGCTATG
cfa2155	5AL	55	F: TTTGTTACAACCCAGGGGG; R: TTGTGTGGCGAAAGAAACAG
cfd8	5DS	60	F: ACCACCGTCATGTCACTGAG; R: GTGAAGACGACAAGACGCAA
gwml69	6AL	55	F: ACCACTGCAGAGAACACATACG; R: GTGCTCTGCTCTAAGTGTGGG
barc345	6BL	55	F: CGCCAGACTGCTAGGATAATACTTT; R: GCGGCTAGTGCTCCCTCATAAT
barcl121	6DL	60	F: GCGAGCAAACTGATCCCAAAAAG; R: TATCGGTGAGTACGCCAAAAACA
cfa2123	7AS	60	F: CGGTCTTTGTTTGCTCTAAACC; R: ACCGGCCATCTATGATGAAG
wmc476	7BS	55	F: TACCAACCACACCTGCGAGT; R: CTAGATGAACCTTCGTGCGG
gwm44	7DS	65	F: GTTGAGCTTTTCAGTTCGGC; R: ACTGGCATCCACTGAGCTG

附表1

图1

图2

表2

图3

图4

图5

附表2

表3

图6

表4

水稻叶片卷曲相关基因在小麦中的同源基因"

水稻基因 Rice gene	基因注释 Gene annotation	部分同源群 Homoeologous groups	小麦的同源基因 Wheat gene
OsACL1	Os04g33860	2	TraesCS2A01G296600 TraesCS2B01G312800 TraesCS2D01G294500
OsDEK1	Os02g47970	6	TraesCS6A01G275500 TraesCS6B01G302900 TraesCS6D01G255800
OsCFL1	Os02g31140	4	TraesCS4A01G407300 TraesCS4B01G306500 TraesCS4D01G304700
OsMYB103L	Os08g05520	7	TraesCS7A01G304500 TraesCS7B01G204800 TraesCS7D01G300000
OsVIL3	Os02g05840	6	TraesCS6A02G123400 TraesCS6B02G151600 TraesCS6D02G113600
OsAGO7	Os03g33650	2	TraesCS2A01G414800 TraesCS2B01G434000 TraesCS2D01G412100
OsCOW1	Os03g06654	4	TraesCS4A01G027500 TraesCS4B01G278300 TraesCS4D01G276600
OsCSLD4	Os12g36890	5	TraesCS5A01G090500 TraesCS5B01G096200 TraesCS5D01G102600
OsHOX32	Os03g43930	5	TraesCS5A01G375800 TraesCS5B01G378000 TraesCS5D01G385300
OsYAB1	Os07g06620	5	TraesCS5A01G371500 TraesCS5B01G373600 TraesCS5D01G380900
OsZHD1	Os09g29130	5	TraesCS5A01G246500 TraesCS5B01G243800 TraesCS5D01G253100
OsPSL1	Os01g19170	1	TraesCS1A01G311100 TraesCS1B01G322500 TraesCS1D01G311000
OsREL1	Os01g64380	2	TraesCS2A01G065100 TraesCS2B01G077300 TraesCS2D01G063200
OsREL2	Os10g41310	1	TraesCS1A01G218200 TraesCS1B01G231600 TraesCS1D01G219800
Osrl14	Os10g40960	1	TraesCS1A01G221200 TraesCS1B01G234500 TraesCS1D01G222800
OsROC5	Os02g45250	6	TraesCS6A01G255800 TraesCS6B01G269700 TraesCS6D01G237000
OsRoc8	Os06g10600	6	TraesCS6A01G324500 TraesCS6B01G354900 TraesCS6D01G304300
OsSLL1	Os09g23200	5	TraesCS5A01G203200 TraesCS5B01G201900 TraesCS5D01G209600
OsSRL2	Os03g19520	4	TraesCS4A01G121600 TraesCS4B01G181700 TraesCS4D01G184300
OsSrl1	Os07g01240	2	TraesCS2A01G256400 TraesCS2B01G286100 TraesCS2D01G347200
OsOFP1	Os01g12690	3	TraesCS3A01G151800 TraesCS3B01G178800 TraesCS3D01G159700
OsMKB3	Os03g52320	4	TraesCS4A01G250600 TraesCS4B01G064000 TraesCS4D01G062900
OsLBD3-7	Os03g57670	5	TraesCS5A01G515300 TraesCS5B01G282700 TraesCS5D01G291300
OsARVL4	Os04g33580	1	TraesCS1A01G036100 TraesCS1B01G231600 TraesCS1D01G219800
OsARF18	Os06g47150	7	TraesCS7A01G446900 TraesCS7B01G346700 TraesCS7D01G436800
OsWOX3A	Os12g01120	5	TraesCS5A01G157300 TraesCS5B01G156400 TraesCS5D01G162600

表4

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标记 Marker	SNP	F₁	F₂	R
RLB	T/C	CCAACCCTAGCTTGTGATCCC	TCCAACCCTAGCTTGTGATCCT	AGGGGCCCACAAGCCTCCTG
RLC	C/G	GAATCACGAAAGCACCCTCGC	GAATCACGAAAGCACCCTCGG	GGCAAACGTGTAGATCGTCTCCC
RLF	A/C	CGTAACATACAACCATGCCTACCT	GTAACATACAACCATGCCTACCG	CGTTCAAGGGTTCACATAAGGCTTTTG
RLH	A/C	GTGGCATGTCAACTCGCGATTTATA	GGCATGTCAACTCGCGATTTATC	CAACTGAGTGGCATATATATAGTCTGTTATA
RLI	A/C	CAGAATCACTTCTTGACTACCCCAA	GAATCACTTCTTGACTACCCCAC	CAGCCTTCTAGAGCCACATTCTAGTA
RLJ	T/C	GTGCTCCTAGGAAGTAGGAGC	ATGTGCTCCTAGGAAGTAGGAGT	CCAAAAGCAAGGACGCTTCCAGG
RLK	T/C	TATTTTGAGACGAAGAGAGAATCGTG	ATTTATTTTGAGACGAAGAGAGAATCGTA	AGGCGGGTGCTTGCCCTTACAT
RLO	C/G	TTATACTAAAGCTGCATCAATTAACTTGC	TTATACTAAAGCTGCATCAATTAACTTGG	GGGCAGTACAACAAACATCAGAAATAAC

性状Trait	种子萌发15 d地上部鲜重 Aboveground fresh weight at 15 days of germination (g)	种子萌发15 d地下部鲜重 Subsurface fresh weight at 15 days of germination (g)	种子萌发15 d 根数目 Root number at 15 days of germination	种子萌发15 d 最大根长 Maximum root length at 15 days of germination (cm)	抽穗期 Heading stage (d)	成熟期 Maturity stage (d)	株高 Plant height (cm)	旗叶面积 Flag leaf area (cm²)	旗叶长 Flag leaf length (cm)	旗叶宽 Flag leaf width (cm)	穗粒数 Grain number per spike	小穗数 Spikelet number	千粒重 1000-grain weight (g)
WT	0.22±0.01^**	0.08±0.01	7±1.3	8.13±0.84	199±1	248±1	78.00±0.28^**	24.62±5.14	21.02±2.65	1.51±0.12^**	56.75±6.65^**	19.25±1.25	52.25±0.91^**
RL1	0.14±0.00	0.08±0.01	7±1.5	8.08±1.26	199±1	247±1	67.67±2.19	19.72±2.51	23.4±2.45	1.09±0.02	35.5±1.91	19.00±0.81	33.70±0.21

杂交组合 Cross combination	F₁表型及卷曲指数 Phenotype of F₁ and leaf rolled index	F₁株数 Number of plants in F₁
RL1×晋麦47 RL1/Jinmai 47	微卷(抽穗期LRIs: 0.64-0.76) Slightly curl (Heading date LRIs: 0.64-0.76)	17
晋麦47×RL1 Jinmai 47/RL1	微卷(抽穗期LRIs: 0.66-0.78) Slightly curl (Heading date LRIs: 0.66-0.78)	18
RL1×临汾5064 RL1/Linfen 5064	微卷(抽穗期LRIs: 0.61-0.76) Slightly curl (Heading date LRIs: 0.61-0.76)	17
临汾5064×RL1 Linfen 5064/RL1	微卷(抽穗期LRIs: 0.68-0.80) Slightly curl (Heading date LRIs: 0.68-0.80)	18

杂交组合 Cross combination	总株数 Number of plants	平展叶 Spread leaf	轻微卷叶 Slightly coiled leaf	高度卷叶 Highly rolled leaf	理论比例 Ratio of theory	卡方 χ²
RL1/晋麦47 RL1/Jinmai 47	176	45	94	37	1:2:1	0.03
RL1/临汾5064 RL1/Linfen 5064	173	44	95	34	1:2:1	0.01

小麦卷叶突变体RL1的生理特性及遗传研究

Physiological characteristics and genetic research of rolled leaf mutant1 (RL1) in wheat (Triticum aestivum L.)

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