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作物学报

作物学报 ›› 2023, Vol. 49 ›› Issue (6): 1584-1600.doi: 10.3724/SP.J.1006.2023.21033

所属专题： 小麦：遗传育种·种质资源·分子遗传学

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

烟农系列小麦高产遗传基础解析

王昊1,2,**(), 孙妮娜1,**(), 王矗1,2, 肖露凝1,2, 肖蓓1,2, 李栋3, 刘洁1, 秦冉4, 吴永振4, 孙晗4, 赵春华4, 李林志1,*(), 崔法4,*(), 刘伟1,*()   

  1. 1山东省烟台市农业科学研究院, 山东烟台 265500
    2烟台大学生命科学学院, 山东烟台 264005
    3山东省种子管理总站, 山东济南 250131
    4鲁东大学农学院/山东省高等学校作物高产抗逆分子模块重点实验室, 山东烟台 264025
  • 收稿日期:2022-04-30 接受日期:2022-10-10 出版日期:2023-06-12 网络出版日期:2022-10-24
  • 通讯作者: *李林志, E-mail: linzhili2002@163.com;崔法, E-mail: sdaucf@126.com;刘伟, E-mail: liuweisdau@163.com
  • 作者简介:王昊，E-mail: whshape@163.com;
    孙妮娜，E-mail: sun200436@163.com第一联系人：**同等贡献
  • 基金资助:
    山东省农业良种工程项目(2019LZGC001);山东省高等学校青创科技支持计划项目(2019KJF002);烟台市科技计划项目(2022XCZX092);财政部和农业农村部国家现代农业产业技术体系建设专项(小麦, CARS-3-2-23);烟台市2022年种子工程攻坚项目——小麦育种攻关联合团队创建项目

Genetic basis analysis of high-yielding in Yannong wheat varieties

WANG Hao1,2,**(), SUN Ni-Na1,**(), WANG Chu1,2, XIAO Lu-Ning1,2, XIAO Bei1,2, LI Dong3, LIU Jie1, QIN Ran4, WU Yong-Zhen4, SUN Han4, ZHAO Chun-Hua4, LI Lin-Zhi1,*(), CUI Fa4,*(), LIU Wei1,*()   

  1. 1Yantai Academy of Agricultural Sciences, Yantai 265500, Shandong, China
    2College of Life Sciences, Yantai University, Yantai 264005, Shandong, China
    3Shandong Provincial Seed Management Station, Jinan 250131, Shandong, China
    4College of Agriculture, Ludong University/Key Laboratory of Crop High Yield and Stress Resistance Molecular Modules in Shandong Higher Education Institutions, Yantai 264025, Shandong, China
  • Received:2022-04-30 Accepted:2022-10-10 Published:2023-06-12 Published online:2022-10-24
  • Contact: *E-mail: linzhili2002@163.com;E-mail: sdaucf@126.com;E-mail: liuweisdau@163.com
  • About author:First author contact:**Contributed equally to this work
  • Supported by:
    Shandong Province Agricultural Improved Seed Engineering Project(2019LZGC001);Shandong Province College Youth Innovation Technology Support Program(2019KJF002);Yantai Science and Technology Plan Project(2022XCZX092);China Agriculture Research System of MOF and MARA(Wheat, CARS-3-2-23);Yantai 2022 Seed Engineering Key Project—Wheat Breeding Joint Team Creation

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摘要：

烟农系列小麦品种具有高产、抗病、广适性等特点, 近几年审定的高产多抗品种烟农1212, 曾多次打破全国冬小麦单产纪录, 鲁麦14已衍生出至少214份小麦新品种, 成为重要的骨干亲本。本研究旨在解析烟农系列小麦高产遗传基础, 明确其高产广适关键染色体区段, 为小麦新品种遗传改良提供理论参考。利用小麦55K SNP芯片对38份烟农系列小麦品种、其部分衍生后代品种和244份育成品种(系)组成的自然群体进行基因型扫描, 并进行了多环境表型鉴定。基因型分析结果表明, 自主选育的17份烟农品种之间的遗传相似系数在0.80~0.99之间, 获得了975个高频率共同选择区段, 区段长度变幅为1.00~75.18 Mb, 其中在2D、4D、6D、7B染色体上存在较多的高频率共同选择区段, 其总长度占对应染色体的40%以上。骨干亲本鲁麦14对其23个衍生后代的平均遗传贡献率为71.45%, 在3个(A、B、D)亚基因组的贡献率分别为69.63%、66.04%和79.82%; 共检测到430个在鲁麦14衍生后代中高频率选择遗传区段, 265个区段(61.6%)与烟农系列高频率共同选择区段重叠。基于自然群体表型鉴定及遗传解析结果显示, 骨干亲本鲁麦14和最新选育的高产广适主推品种烟农1212均富集了千粒重和单株产量优异等位基因; 在鲁麦14高频率选择区段上富集了92.3%和84.4%的来自鲁麦14的增加千粒重和单株产量显著关联SNP位点, 主要分布在2A、2B、2D、4A、5B、6A、7A染色体上。烟农系列小麦高频率共同选择基因组区段已富集了丰富的高产基因位点, 是其高产稳产的遗传基础所在。

关键词: 小麦, 遗传贡献, 55K SNP芯片, 遗传区段, 骨干亲本

Abstract:

The Yannong series wheat has the characteristics of high yield potential, disease resistance, and wide adaptability. Yannong 1212, a high-yield and multi-resistant variety approved in recent years, has repeatedly broken the national winter wheat yield record for many times. To date, the numerous varieties derived from Yannong series have been released. Lumai 14 has derived from at least 214 wheat authorized varieties and become an important backbone parent by 2020. This objective of the study is to characterize genetic basis of high yielding potential in Yannong series, identify the key chromosomal segments of high yield and wide adaptability, and provide theoretical reference for the genetic improvement of new wheat varieties. The genotype of 38 Yannong series wheat varieties, some of their derived varieties, and 244 authorized varieties (advanced lines) were scanned by wheat 55K SNP array, and the environmental phenotypes were identified. Phenotype evaluation of yield-related traits of the natural mapping population was conducted in multiple environments. The genetic effects of the common high-frequency co-selected genetic segment (HFCS) of Yannong series varieties were characterized in detail based on the genotypes of wheat 55K SNP array. In addition, the HFCS from Lumai 14 to its derived varieties were also specified. The genetic basis of high yield potential in Yannong 1212 and Lumai 14 were revealed at the whole genome-wide level. The genetic similarity coefficients among the 17 Yannong series authorized varieties ranged from 0.80 to 0.99. Based on genotyping data from the 55K SNP array in the 17 Yannong series authorized varieties, 975 HFCS were obtained, and their segments lengths ranged from 1.00 Mb to 75.18 Mb. Most HFCS were distributed on chromosomes 2D, 4D, 6D, and 7B, with the total length accounting for more than 40% of the corresponding chromosomes, respectively. The genetic contribution rate of Lumai 14 to its 23 derived varieties was approximately 71.45% on average at the genome level, with A, B, and D of 69.63%, 66.04%, and 79.82%, respectively. A total of 430 high-frequency transmission genetic segments (HFTGS) 14 were detected in Lumai derived progency and 265 blocks (61.6%) overlapped with the HFSC in the Yannong series. Lumai 14 was a backbone parents and Yannong 1212 was a novel authorized varieties with high yield potential and wide adaptability. To characterize their genetic basis of high yield and wide adaptability, single marker analysis based on the natural population was performed. The results showed that both Lumai 14 and Yannong 1212 had been enriched in excellent alleles for thousand kernel weight (TKW) and yield per plant (YPP). Approximately 92.3% and 84.4% of the significant loci in the HFTGS of Lumai 14 were shown to increase TKW and YPP, mainly distributing on chromosomes 2A, 2B, 2D, 4A, 5B, 6A, and 7A. Yannong series authorized wheat varieties had enriched in excellent alleles of yield-related genes and QTLs especially for TKW and YPP, which played a key role to their high and stable yielding potential.

Key words: wheat, genetic contribution, 55K SNP array, genetic segment, backbone parent

表1

试验材料名称、组合、审(认)定年份"

编号
Serial		 品种
Variety		 组合
Combination		 审/认定年份
Approved and identified year
1 烟农15 Yannong 15 蚰包//意大利ST2422/464 Youbao//Italy ST2422/464 1982
2 鲁麦14 Lumai 14 C149/F4530 1990, 1992, 1993
3 鲁麦21 Lumai 21 鲁麦13/宝丰7228 Lumai 13/Baofeng 7228 1996
4 烟农22 Yannong 22 鲁麦14//尉132/87初20 Lumai 14//Wei 132/87 Chu 20 2002
5 济麦20 Jimai 20 鲁麦14/884187 Lumai 14/884187 2003, 2004
6 烟农23 Yannong 23 烟1061/鲁麦14 Yan 1061/Lumai 14 2003
7 良星99 Liangxing 99 91102/鲁麦14//PH85-16 91102/Lumai 14//PH85-16 2004, 2006
8 泰山22 Taishan 22 鲁麦18/鲁麦14 Lumai 18/Lumai 14 2004
9 烟农24 Yannong 24 陕229/安麦1号 Shaanxi 229/Anmai 1 2004
10 济麦22 Jimai 22 935024/935106 2006, 2007
11 青丰1号Qingfeng 1 鲁麦14/烟农15 Lumai 14/Yannong 15 2006
12 烟2415 Yan 2415 烟849/鲁麦21 Yan 849/Lumai 21 2006
13 烟农5158 Yannong 5158 烟航选2号/烟农15 Yanhangxuan 2/Yannong 15 2007, 2009, 2010
14 泰农18 Tainong 18 莱州137/烟369-7 Laizhou 137/Yan 369-7 2008
15 山农17 Shannong 17 L156/莱州137 L156/Laizhou 137 2009
16 青农2号 Qingfeng 2 鲁麦14/烟农15//矮秆麦
Lumai 14/Yannong 15//Dwarf wheat		 2010
17 泰农19 Tainong 19 莱州137/济南17 Laizhou 137/Jinan 17 2011
18 烟农999 Yannong 999 烟航选2号/临9511//烟BLU14-15
Yanhangxuan 2/Lin 9511//Yan BLU14-15		 2011, 2016, 2018
19 石农086 Shinong 086 鲁麦14/邯6172 Lumai 14/Han 6172 2014
20 烟农836 Yannong 836 山农721511/鲁麦21 Shannong 721511/Lumai 21 2010, 2014
21 济麦23 Jimai 23 豫麦34/济麦22 Yumai 34/Jimai 22 2016
22 烟农173 Yannong 173 济麦22/烟2415 Jimai 22/Yan 2415 2016
23 冀麦738 Jimai738 藁9618/良星99 Gao 9618/Liangxing 99 2016
24 石麦25 Shimai 25 济麦22/金禾9123 Jimai 22/Jinhe 9123 2016
25 中信麦99 Zhongxinmai 99 良星99/222 Liangxing 99/222 2016
编号
Serial		 品种
Variety		 组合
Combination		 审/认定年份
Approved and identified year
26 轮选266 Lunxuan 266 济麦19/济麦22//济麦22 Jimai 19/Jimai 22//Jimai 22 2018
27 泰科麦31 Taikemai 31 泰山26/淮麦20 Taishan 26/Huaimai 20 2018
28 淄麦29 Zimai 29 泰农18//烟5072 Tainong 18//Yan 5072 2018
29 邯麦19 Hanmai 19 邯02-6018/济麦22 Han 02-6018/Jimai 22 2018
30 中麦23 Zhongmai 23 济麦22/淮9701 Jimai 22/Huai 9701 2019
31 烟农1212 Yannong 1212 烟5072/石94-5300 Yan 5072/Shi 94-5300 2018, 2019, 2021
32 烟农377 Yannong 377 By114/烟6089 By114/Yan 6089 2020
33 烟农215 Yannong 215 烟672/烟农999 Yan 672/Yannong 999 2020
34 烟农161 Yannong 161 济麦22/烟农1212 Jimai 22/Yannong 1212 2021
35 烟农301 Yannong 301 济麦22/烟1201 Jimai 22/Yan 1201 2021
36 烟农1766 Yannong 1766 烟农09135 /济麦22 Yannong 09135/Jimai 22 区试1年
Regional test varieties for one year
37 烟农30 Yannong 30 烟农09135 /济麦22 Yannong 09135/Jimai 22 区试2年
Regional test varieties for two years
38 烟农31 Yannong 31 烟农09135 /济麦22 Yannong 09135/Jimai 22 区试1年
Regional test varieties for one year

图1

烟农系列小麦品种的UPGMA聚类图"

表2

不同染色体的高频率共同选择遗传区段分布情况"

染色体
Chromosome		 区段数
Number of segments		 区段总长
Total segments length (Mb)		 染色体总长
Chromosome length (Mb)		 区段占比
Segments’ proportion (%)
1A 57 219.93 602 36.53
1B 42 267.54 752 35.58
1D 19 65.61 503 13.04
2A 89 292.47 794 36.84
2B 69 317.71 800 39.71
2D 30 305.46 654 46.71
3A 22 81.68 762 10.72
3B 58 191.00 845 22.60
3D 28 83.15 616 13.50
4A 72 194.43 753 25.82
4B 25 133.95 675 19.85
4D 72 213.40 519 41.12
5A 37 257.47 718 35.86
5B 33 133.51 716 18.65
5D 19 53.17 572 9.30
6A 34 237.01 618 38.35
6B 73 241.14 712 33.87
6D 40 200.17 490 40.85
7A 62 321.62 744 43.23
7B 59 356.53 762 46.79
7D 35 90.63 648 13.99

表3

全基因组水平鲁麦14对其23个衍生品种遗传贡献率"

衍生材料
Derived varieties		 A基因组
A genome		 B基因组
B genome		 D基因组
D genome		 全基因组
Whole genome
济麦20 Jimai 20 68.25 65.57 78.66 70.32
济麦22 Jimai 22 69.41 63.33 80.57 70.75
济麦23 Jimai 23 68.70 64.98 81.28 71.07
良星99 Liangxing 99 69.43 69.75 80.83 72.58
轮选266 Lunxuan 266 69.41 63.34 80.67 70.78
青丰1号 Qingfeng 1 70.88 66.77 81.33 72.57
青农2号 Qingnong 2 70.85 66.67 81.43 72.55
山农17 Shannong 17 69.11 66.02 81.08 72.84
泰科麦31 Taikemai 31 70.38 64.23 81.28 71.63
泰农18 Tainong 18 69.08 63.19 80.62 70.58
泰农19 Tainong 19 69.08 63.29 80.67 70.62
泰山22 Taishan 22 69.78 66.32 77.35 70.88
烟农173 Yannong 173 69.08 65.03 79.62 70.81
烟农22 Yannong 22 80.92 69.25 80.17 77.54
烟农23 Yannong 23 68.96 66.32 81.03 71.48
烟农999 Yannong 999 66.78 66.22 80.12 70.21
中麦23 Zhongmai 23 68.85 64.18 79.47 70.43
淄麦29 Zimai 29 69.32 67.81 80.62 71.94
邯麦19 Hanmai 19 69.11 66.02 74.23 69.65
冀麦738 Jimai 738 70.29 68.06 82.49 72.96
石麦25 Shimai 25 66.22 69.60 79.97 70.84
石农086 Shinong 086 68.73 69.25 72.47 69.88
中信麦99 Zhongxinmai 99 68.88 63.64 79.72 70.36
平均值 Average 69.63 66.04 79.81 71.45

图2

产量相关的QTL在高频率共同选择遗传区段上的分布"

图3

小麦骨干亲本鲁麦14及其部分衍生品种的UPGMA聚类图"

图4

小麦骨干亲本鲁麦14及其部分衍生品种的系谱关系图"

表4

鲁麦14高频率共同选择遗传区段在不同染色体分布"

染色体
Chromosome		 区段数
Number of segments		 区段总长
Total segments length (Mb)		 染色体总长
Chromosome length (Mb)		 区段占比
Segments’ proportion (%)
1A 37 95.88 602 15.93
1B 9 26.48 752 3.52
1D 5 9.16 503 1.82
2A 35 94.48 794 11.90
2B 23 69.23 800 8.65
2D 19 58.96 654 9.01
3A 8 21.91 762 2.88
3B 17 41.93 845 4.96
3D 5 12.19 616 1.98
4A 77 186.03 753 24.71
4B 9 36.98 675 5.48
4D 18 69.80 519 13.45
5A 11 26.67 718 3.71
5B 13 23.14 716 3.23
5D 6 17.98 572 3.14
6A 41 103.25 618 16.71
6B 18 35.33 712 4.96
6D 21 52.36 490 10.69
7A 32 85.46 744 11.49
7B 18 57.44 762 7.54
7D 8 61.76 648 9.53

图5

基于鲁麦14高频率选择区段(位点)在产量性状上的增效作用解析 每个单元分为3个部分, 左侧绿色为染色体高频率选择区段, 中间为区段LOD值热图, 右侧为区段内产量性状的增效作用值, 包括红、黄、蓝、紫4个部分, 分别代表千粒重、单株穗数、穗粒数、单株产量。"

图6

基于鲁麦14和烟农1212 基因型在产量性状上增效位点占比 A: 鲁麦14产量相关性状增效SNP位点占比; B: 烟农1212产量相关性状增效SNP位点占比; 其中蓝色部分代表增效位点占比, TKW:千粒重; SNPP: 单株穗数; KNS: 穗粒数; YPP: 单株产量。"

附表1

鲁麦14高频率共选区段中的标记信息"

染色体
Chromosome		 高频率共选区段
HFCS		 标记
Markers		 关联性状
Associated Traits
1D 58.905983-60.704201 Xgwm458, wPt-741323--wPt-664609 seed weight from sample ears, spike number
2A 112.911673-122.169547 Xgwm356, Xgwm526.1, Xgwm265, 1128252—985175, Xgwm356--Xgwm382 thousand kernel weight, spike number per plant, tiller number, grain yield, spike length
125.065333-130.287176 Xgwm382 kernel weight
2B 80.919673-81.943795 Xgwm120 grain yield
3A 46.256526-49.846165 wPt-7756 spike length
112.292542-114.888625 wpt-1888 kernel number per spike
3B 39.830393-40.866704 Tdurum_contig50954_1393--Kukri_c15654_309 grain Weight
96.503834-103.237886 wPt-8752--wPt-1171, wPt-1804, wPt-4933--wPt-667746 grain number per spike, spike length, spikelet number per spike, grain yield
4A 46.408021-53.671828 Xmwg549--Xwg622, WPT-5694-WPT-7939, Xwmc468--Xbarc170, Ku_c6779_1381--Xwmc258 grain number, thousand kernel weight, grain number per spike
57.494839-59.99089 Xmwg549--Xbcd1670, BS00066891_51--wsnp_Ex_c3988_7221220, RAC875_c83401_115--Tdurum_contig65718_209, wPt-2247--tPt-9400, tPt-9400, Xwmc258, 1242399--3064552 grain number, grain weight per ear, spike length, kernels per spike, number of nodes, kernels per node, fertile spikelet number, thousand grain weight
65.519761-66.770139 IWB25909, wPt-0023, wPt-4620, wsnp_Ku_c9746_16265584--BS00087277_51 kernel weight, thousand grain weight, grain yield, spikelet number per spike
75.307817-83.610777 Xcdo545, Xgwm160--Xwmc232, Xcfd2 yield, grain number per spike, spike length, compactness, tiller number
83.759753-85.587069 Xbarc78--Xwmc722 sterile spikelet number
88.019734-89.066785 Xwmc497--Xwmc219 grain number per spike
4D 57.867989-62.949715 wmc720, Xbarc48--Xbarc240 thousand grain weight
5A 50.743753-53.276562 Tdurum_contig86202_175--wsnp_Ra_c10915_17838202, Xbarc141, Xgwm415--Xgwm304, wsnp_BE404341A_Ta_2_3, wPt-4262--VRNA1, Xgwm293 flowering, thousand grain weight, grain number per ear, spike length, grain yield, grain weight per plant, spike number per plant, grain weight per ear, grain number per total spikelets, spikelet number per spike, grain number per square meter
5B 33.477304-34.80488 wsnp_Ex_c49423_54028488--wsnp_Ra_c39562_47242455 spikelet number per spike
39.416962-41.823352 xcfd7, Xbarc340.1--Xgwm443.2, wPt-6014--wPt-7006 grain filling rate, grain number per spike, kernels per node
89.684338-93.239903 Xgwm371--Xgwm604 yield, thousand kernel weight
6B 62.268859-66.443909 WPT-669607, wPt-1048--Xgwm219, XksuG30--Xfbb169, Xbarc178 kernel weight, thousand kernel weight, grain weight per ear
78.17482-79.349348 IWB5488, wPt-1264 spike length, kernels per node
80.980565-82.649349 wPt-6116--wPt-1541, wPt-0171, RAC875_c28848_330, Tdurum_contig68258_1773 penetrance of clavate architecture, kernels per spikelet, grain number per spike, grain weight
6D 29.901228-31.937716 Xgdm132 grain yield
34.387538-38.576284 Xgwm469 spike number per plant
67.499205-68.923987 xbarc196--xgwm325 grain filling rate
102.752092-106.435316 Xbcd1319--XksuD27 biomass, grain yield, tiller number
7A 101.609097-108.885105 Xgwm332, Kukri_rep_c97425_164, IWA7325--IWA4626, 2262955, gwm282, IWA7325--IWA4626, 2262955--Xwmc633 grain yield, tiller number, grain number per ear, thousand kernel weight, grain number per spike
7B 57.805476-60.081873 Xgwm46, wsnp_BE404339B_Ta_2_2--BS00059061_51, WPT-4230-wmc517, Xwmc758--Xbarc72 thousand kernel weight, days to heading, flowering date, spikelet number per spike
67.969371-68.990187 Xpsr927, Xwmc396 grain weight/plant, grain weight/ear, grain number per square meter
71.476476-73.787477 Xbarc50 spike length, floret number per spikelet, grain number per spike
136.182314-137.23701 Xmwg710a, Xwmc526--Xwmc273 final grain weight, tiller number
7D 96.302557-101.231804 Xbarc26, xgwm437--xcfd14, xgwm437, Xcfd14--Xbarc172 thousand grain weight, biomass, spike weight, grain number per spike
137.311934-142.701669 Xgwm428 grain number per ear
154.122529-166.657341 Xwmc634--Xwmc273, Xwmc634, Xbarc76, Xcfd69, Xgwm37 fertile spikelet number, spike number, grain number per spike, grain yield, thousand kernel weight
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