Phenotypic Analysis and Preliminary Mapping of Wheat Dwarf Small⁃Kernel Mutant smk56

doi:10.15933/j.cnki.1004-3268.2025.02.003

Abstract

Abstract: A dwarf and small grain mutant smk56（small kernel 56）with stable phenotypic inheritance was obtained from the seeds of wheat variety Zhoumai 22，which was mutagenized by China’s first microgravity science experiment satellite，Shijian⁃10.The phenotype and endosperm starch morphology of smk56 were analyzed and observed.The mutant was hybridized with the wild type Zhoumai 22（WT）to construct an F₂segregating population.The plant height，thousand⁃grain weight and other agronomic traits of F_2：3 families were investigated，and the genetic analysis of the mutant gene was carried out.Then，the bulked segregant analysis sequencing（BSA⁃Seq），exon capture sequencing and linkage analysis were used for preliminary location of the mutant gene.The results showed that compared with WT，smk56 had no significant difference in seedling habits，leaf color，panicle type，heading stage and mature stage；Plant height of smk56 significantly decreased by 14.51%，and thousand⁃grain weight significantly decreased by 21.83%；Additionally，spike length，effective spikelet number，internode length below the spike，grain number per spike，grain length and grain width were all significantly or extremely significantly decreased；The starch granules of smk56 had poor filling degree and a more loosely arrangement structure，with underdeveloped starch bodies in endosperm cells，but there was no significant difference in the arrangement and shape regularity of starch granules between smk56 and WT.Genetic analysis indicated that the smk56 trait was controlled by a single recessive gene，which was preliminarily mapped to a region on the wheat chromosome 6D，spanning from 23.18 cM to 35.54 cM on the genetic map，and closely linked to the marker gwm469，with the flanking molecular markers of cfd1 and cfd132.In summary，the dwarf and small grains of wheat mutant smk56 may be caused by the shortening of stem internodes and the poor development of amyloplasts in grain endosperm cells；The target gene is preliminarily mapped to the wheat chromosome 6D genetic map of 23.18—35.54 cM，with a genetic distance of 12.36 cM，and it is speculated that this region may habor a novel important gene related to the formation of plant height and thousand⁃grain weight in wheat.

Key words: Wheat, Plant height, Thousand?grain weight, smk56, Genetic analysis, Gene mapping

摘要： 利用我国首颗微重力科学试验卫星“实践十号”航天诱变小麦品种周麦22获得表型稳定的矮秆小粒突变体smk56（Small kernel 56），对其表型和胚乳淀粉形态进行观察，并将smk56与野生型周麦22（WT）杂交获得F₂分离群体，调查F_2：3家系的株高和千粒质量，进行突变基因遗传分析，然后通过混合群体分离测序（BSA-Seq）技术、外显子捕获测序和连锁分析进行基因初步定位研究。结果表明，与WT相比，突变体smk56幼苗习性、叶色、穗型、抽穗期和成熟期均无显著差异，株高极显著降低14.51%，千粒质量极显著降低21.83%，穗长、有效小穗数、穗下节长、穗粒数、粒长和粒宽均显著或极显著降低；smk56的淀粉颗粒间充实度差，排列较疏松，胚乳细胞淀粉体发育差，而淀粉颗粒排列和形状与WT相比无明显差异。遗传分析结果表明，smk56突变性状受隐性单基因控制，基因突变位点初步定位于小麦6D染色体遗传图谱23.18—35.54 cM，且与gwm469标记紧密连锁，侧翼分子标记分别为cfd1与cfd132。综上，小麦突变体smk56 的矮秆小粒表型可能由茎节间缩短和籽粒胚乳细胞淀粉体发育差所致，控制该性状的目标基因定位于6D染色体遗传图谱23.18—35.54 cM，遗传距离为12.36 cM，推测该区段可能存在一个新的控制小麦株高和千粒质量形成相关的重要基因。

关键词: 小麦, 株高, 千粒质量, smk56, 遗传分析, 基因定位

CLC Number:

S512

ZHANG Fuyan, SUN Congwei, CHEN Xiaojie, WANG Jiahuan, CHENG Zhongjie, ZHAO Wan, MA Xuhui, ZHANG Jianwei, FAN Jialin. Phenotypic Analysis and Preliminary Mapping of Wheat Dwarf Small⁃Kernel Mutant smk56[J]. Journal of Henan Agricultural Sciences, 2025, 54(2): 21-29.

张福彦, 孙丛苇, 陈晓杰, 王嘉欢, 程仲杰, 赵婉, 马旭辉, 张建伟, 范家霖. 小麦矮秆小粒突变体smk56表型分析与基因初步定位[J]. 河南农业科学, 2025, 54(2): 21-29.

Figures/Tables 6

References

［1］蒋赟，张丽丽，薛平，等. 我国小麦产业发展情况及国际经验借鉴［J］. 中国农业科技导报，2021，23（7）：1⁃10.
JIANG Y，ZHANG L L，XUE P，et al. Development status of wheat industry in China and international experience for reference［J］.Journal of Agricultural Science and Technology，2021，23（7）：1⁃10. ［2］茹振钢，冯素伟，李淦. 黄淮麦区小麦品种的高产潜力与实现途径［J］.中国农业科学，2015，48（17）：3388⁃3393.
RU Z G，FENG S W，LI G. High⁃yield potential and effective ways of wheat in Yellow & Huai River valley facultative winter wheat region［J］.Scientia Agricultura Sinica，2015，48（17）：3388⁃3393.

［3］ZHANG S Y，ZHANG X H，QIU X L，et al. Quantifying the spatial variation in the potential productivity and yield gap of winter wheat in China［J］.Journal of Integrative Agriculture，2017，16（4）：845⁃857.

［4］罗江陶，郑建敏，邓清燕，等.2000—2020年四川小麦育成品种产量增益分析［J］.中国农业科学，2024，57（20）：3945⁃3956.

LUO J T，ZHENG J M，DENG Q Y，et al.Yield gain analysis of wheat varieties in Sichuan from 2000 to 2020［J］.Scientia Agricultura Sinica，2024，57（20）：3945⁃3956.

［5］周晓变，赵磊，陈建辉，等.黄淮麦区小麦种质资源矮秆基因分布及其与农艺性状的关系［J］.麦类作物学报，2017，37（8）：997⁃1007.

ZHOU X B，ZHAO L，CHEN J H，et al.Distribution of dwarf genes and their association with agronomic traits in bread wheat from the yellow and Huai wheat region［J］.Journal of Triticeae Crops，2017，37（8）：997⁃1007.

［6］MO Y J，VANZETTI L S，HALE I，et al.Identification and characterization of Rht25，a locus on chromosome arm 6AS affecting wheat plant height，heading time，and spike development［J］.Theoretical Applied Genetics，2018，131（10）：2021⁃2035.

［7］SUN L H，YANG W L，LI Y F，et al.A wheat dominant dwarfing line with Rht12，which reduces stem cell length and affects gibberellic acid synthesis，is a 5AL terminal deletion line［J］. The Plant Journal，2019，97（5）：887⁃900.

［8］ZHAO K，XIAO J，LIU Y，et al. Rht23（5Dq'）likely encodes a Q homeologue with pleiotropic effects on plant height and spike compactness［J］.Theoretical and Applied Genetics，2018，131：1825⁃1834.

［9］PEARCE S，SAVILLE R，VAUGHAN S P，et al.Molecular characterization of Rht⁃1 dwarfing genes in hexaploid wheat［J］.Plant Physiology，2011，157（4）:1820⁃1831.

［10］XIONG H C，ZHOU C Y，FU M Y，et al.Cloning and functional characterization of Rht8，a ‘Green Revolution’replacement gene in wheat［J］.Molecular Plant，2022，15（3）：373⁃376

［11］CHAI L L，XIN M M，DONG C Q，et al.A natural variation in ribonuclease H⁃like gene underlies Rht8 to confer ‘Green Revolution’ trait in wheat［J］.Molecular Plant，2022，15（3）：377⁃380.

［12］WU W，LI C J，MA B L，et al. Genetic progress in wheat yield and associated traits in China since 1945 and future prospects［J］. Euphytica，2014，196（2）：155⁃168.

［13］魏玮，郭嘉莲，万琳涛，等. 小麦粒重形成的分子调控机制研究综述［J］.浙江农林大学学报，2016，33（2）：348⁃356.

WEI W，GUO J L，WAN L T，et al.Research progress on molecular regulation mechanism of grain weight formation in wheat［J］.Journal of Zhejiang A & F University，2016，33（2）：348⁃356.

［14］YANG J，ZHOU Y J，WU Q H，et al.Molecular characterization of a novel TaGL3‑5A allele and its association with grain length in wheat（Triticum aestivum L.）［J］.Theoretical and Applied Genetics，2019，132（6）：1799⁃1814.

［15］NIAZ M，ZHANG L R，LV G G，et al. Identification of TaGL1‑B1 gene controlling grain length through regulation of jasmonic acid in common wheat［J］. Plant Biotechnology Journal，2023，21（5）：979⁃989.

［16］WANG S S，ZHANG X F，CHEN F，et al. A single⁃nucleotide polymorphism of TaGS5 gene revealed its association with kernel weight in Chinese bread wheat［J］. Frontiers in Plant Science，2015，6：1166.

［17］YAN X F，ZHAO L，REN Y，et al.Genome⁃wide association study revealed that the TaGW8 gene was associated with kernel size in Chinese bread wheat［J］.Scientific Reports，2019，9（1）：2702.

［18］MA D Y，YAN J，HE Z H，et al.Characterization of a cell wall invertase gene TaCwi‑A1 on common wheat chromosome 2A and development of functional markers ［J］. Molecular Breeding，2012，29（1）：43⁃52.

［19］MA M，ZHAO H X，LI Z J，et al.TaCYP78A5 regulates seed size in wheat（Triticum aestivum）［J］.Journal of Experimental Botany，2016，67（5）：1397⁃1410.

［20］VOLPICELLA M，FANIZZA I，LEONI C，et al.Identification and characterization of the sucrose synthase 2 gene（Sus2）in durum wheat［J］.Frontiers in Plant Science，2016，7：266.

［21］张福彦，范家霖，陈晓杰，等. 小麦粒重相关基因的遗传定位和分子标记辅助育种进展［J］.植物遗传资源学报，2020，21（3）：507⁃516.

ZHANG F Y，FAN J L，CHEN X J，et al. Genetic localization and marker assisted breeding of grain weight⁃related genes in common wheat［J］. Journal of Plant Genetic Resources，2020，21（3）：507⁃516.

［22］吕广德，靳雪梅，郭营，等.小麦株高分子遗传学研究进展［J］.植物遗传资源学报，2021，22（3）：571⁃582.

LÜ G D，JIN X M，GUO Y，et al.Advances in molecular genetics of wheat plant height［J］. Journal of Plant Genetic Resources，2021，22（3）：571⁃582.

［23］ZOU C，WANG P X，XU Y B. Bulked sample analysis in genetics，genomics and crop improvement［J］.Plant Biotechnology Journal，2016，14（10）：1941⁃1955.

［24］卞能飞，温明星，高润红，等.普通小麦望水白矮秆多蘖突变体的鉴定及遗传分析［C］//中国遗传学会. 中国遗传学会第九次全国会员代表大会暨学术研讨会论文集. 哈尔滨：中国遗传学会，2013：128.

BIAN N F，WEN M X，GAO R H，et al.Identification and genetic analysis of dwarf and tiller mutant of common wheat Wangshuibai［C］// Genetics Society of China.Proceeding of The 9th National Congress and Academic Seminar of Genetics Society of China.Harbin：Genetics Society of China，2013：87.

［25］付美玉，熊宏春，周春云，等.小麦矮秆突变体je0098的遗传分析与其矮秆基因定位［J］.作物学报，2022，48（3）：580⁃589.

FU M Y，XIONG H C，ZHOU C Y，et al.Genetic analysis of wheat dwarf mutant je0098 and molecular mapping of dwarfing gene［J］.Acta Agronomica Sinica，2022，48（3）：580⁃589.

［26］WANG F X，YU Z P，ZHANG M L，et al.ZmTE1 promotes plant height by regulating intercalary meristem formation and internode cell elongation in maize［J］.Plant Biotechnology Journal，2022，20（3）：526⁃537.

［27］金久岩，姜文洙，严永峰，等. 水稻矮秆小粒突变体dsg-j的鉴定及基因定位［J］.东北农业科学，2024，49（2）：11⁃16.

JIN J Y，JIANG W Z，YAN Y F，et al.Identification and gene mapping of the dwarf small⁃grain mutant dsg‑j in rice［J］.Journal of Northeast Agricultural Sciences，2024，49（2）：11⁃16.

［28］冉莉萍，夏敏洁，姚慧慧，等. 小麦乙烯响应转录因子基因wfzp突变体颖果发育的研究［J］.麦类作物学报，2021，41（12）：1459⁃1468.

RAN L P，XIA M J，YAO H H，et al.Caryopsis development of wfzp mutants of ethylene response transcription factor gene in wheat［J］.Journal of Triticeae Crops，2021，41（12）：1459⁃1468.

［29］何羽喆，徐善斌，陈彦宇，等.水稻糯性突变体r162的鉴定与基因定位［J］.浙江大学学报（农业与生命科学版），2024，50（3）：368⁃381.

HE Y Z，XU S B，CHEN Y Y，et al.Identification and gene mapping of rice glutinous mutant r162［J］.Journal of Zhejiang University（Agriculture and Life Sciences），2024，50（3）：368⁃381.

［30］杜启迪，郭会君，熊宏春，等.小麦顶端小穗退化突变体asd1 基因定位［J］.作物学报，2022，48（8）：1905⁃1913.

DU Q D，GUO H J，XIONG H C，et al.Gene mapping of apical spikelet degeneration mutant asd1 in wheat［J］. Acta Agronomica Sinica，2022，48（8）：1905⁃1913.
［31］ZHANG G Z，WANG Y Y，GUO Y，et al.Characterization and mapping of QTLs on chromosome 2D for grain size and yield traits using a mutant line induced by EMS in wheat［J］.The Crop Journal，2015，3（2）：135⁃144.

［32］陶怡然，熊毓贞，谢佳，等.水稻矮化大粒突变体sdb1的鉴定与基因定位［J］.作物学报，2018，44（11）：1621⁃1630.

TAO Y R，XIONG Y Z，XIE J，et al.Identification and gene mapping of sdb1 mutant with a semi⁃dwarfism and bigger seed in rice［J］.Acta Agronomica Sinica，2018，44（11）：1621⁃1630.
［33］NADOLSKA⁃ORCZYK A，RAJCHEL I K，ORCZYK W，et al. Major genes determining yield⁃related traits in wheat and barley［J］.Theoretical and Applied Genetics，2017，130（6）：1081⁃1098.

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