基于NCII遗传交配设计的籼稻抽穗期全基因组关联分析
徐凯, 郑兴飞, 张红燕, 胡中立, 宁子岚, 李兰芝
作物学报
2023, 49 ( 1):
86-96.
DOI: 10.3724/SP.J.1006.2023.12079
抽穗期受光温资源和基因网络的共同调控, 影响作物产量和品种的地域适应性, 通过关联分析鉴定与抽穗期性状相关的显著位点和基因, 对解析抽穗期的遗传基础具有重要意义。本研究按照双因素交叉式遗传设计(North Carolina design II, NCII)将115份籼稻品种作为父本, 5份不育系作为母本, 进行测交, 得到了575个F1代的测交群体。对亲本和F1代抽穗期的表型值、亲本品种群体的一般配合力、F1代群体的特殊配合力和超亲优势值进行全基因组关联分析, (1) 共定位到104个关联位点, 分布在12条染色体上。其中, 第4条染色体上检测到的显著位点最多, 为16个。在亲本的抽穗期表型、一般配合力、F1代的抽穗期表型、特殊配合力、超亲优势值5个数据集中分别检测到6、5、15、57和21个。对这5个数据集中的显著关联位点进行表型变异分析, 发现在亲本抽穗期表型、一般配合力、F1代抽穗期表型、特殊配合力和超亲优势值这5个数据集中的显著关联位点对表型变异的总贡献率(phenotypic variation explained, PVE)分别为79.57%、10.51%、33.35%、56.42%和54.86%。其中, 25个位点在多个数据集中被检测到, 可能为抽穗期相关的热点区域。(2) 通过关联分析得到的显著位点与日本晴参考基因组注释信息比对, 共检测到5个已克隆抽穗期基因, 其中3个基因与显著关联位点的基因组距离小于200 kb, 对这3个基因中的单倍型组合与单个基因的优异单倍型进行比较发现, 亲本品种群体中单倍型组合SDG724 (Hap.A)_Hd17 (Hap. E)_Ghd7 (Hap. A)的对应的水稻单株籽粒产量较高, 抽穗期较长, 该组合中各基因的单倍型对应于单个克隆基因的优异单倍型, 表明优异单倍型的聚合的常规稻, 具有更高的产量和更长的抽穗期。测交子代未见此情形, 测交子代群体中SDG724 (Hap. I)_Hd17 (Hap. K)_Ghd7 (Hap. I)的单倍型组合形式的水稻有适中的抽穗期和较高的产量, 说明测交子代的抽穗期遗传机制较父本(常规水稻品种)复杂。全基因组关联分析和单倍型分析相结合, 能利用到多个SNP提供的连锁不平衡信息, 提高了基因检测效率, 对培育高产的水稻品种具有重要的指导意义。
数据集 DataSet | 染色体 Chr. | 起始位置 Start | 终止位置 End | −log10 (P)最高的位点 Highest association site | P值负对数 −log10 (P) | 表型方差解释率 Phenotypic variation explained (%) | 克隆基因 Cloned gene | V | 3 | 34355029 | 34355029 | chr03_34355029 | 7.58 | 10.46 | SNAC1; SPIN1; OsPHY2; OsIDS1; gh1; ZFP182; ZFP15; OsADF3; OsRLCK118 | 4 | 4302632 | 4302632 | chr04_4302632 | 5.32 | 24.26 | OsEMF2a; ART2; OsCAS | 6 | 4730505 | 4730505 | chr06_4730505 | 10.14 | 8.89 | OsPTF1; OsERF71; OsKASI; OsARF16 | 9 | 7869103 | 7869103 | chr09_7869103 | 6.76 | 11.15 | OsbZIP71; OsMED25; OsEMF2b; SDG724 | 10 | 16080525 | 16080525 | chr10_16080525 | 6.55 | 22.07 | OsCDC48E; OsABCG25; OsPht1; OsPht1; OsHCI1; OsMTA4 | 12 | 15155333 | 15155333 | chr12_15155333 | 6.76 | 2.73 | PIOX | GCA | 1 | 33603983 | 33603983 | chr01_33603983 | 6.86 | 0.40 | OsGH3-1; OsPME1; OsERF3 | 2 | 4065908 | 4065908 | chr02_4065908 | 5.98 | 5.44 | OsALDH5F1; OsDPK2 | 4 | 20338147 | 20338147 | chr04_20338147 | 5.19 | 0.53 | OSINV4; GIF1; ACL1; COPT4 | 6 | 4745474 | 4745474 | chr06_4745474 | 8.93 | 4.10 | OsPTF1; OsERF71; SUS2; OsKASI; OsARF16; | 11 | 27790672 | 27790672 | chr11_27790672 | 5.62 | 0.04 | JAmyb; Pikp_2 | TC | 1 | 35427407 | 35427407 | chr01_35427407 | 7.16 | 5.07 | OsWRKY24; OsYSL18; LAX1; OsBAG4; OsAMT2 | 2 | 28380703 | 28380703 | chr02_28380703 | 7.16 | 9.44 | OsPIE1; 709 | 3 | 2085577 | 2085577 | chr03_2085577 | 5.39 | 2.61 | RAI1; OsCYP96B4; OsDSS1; sd37; bsh1 | 3 | 14946317 | 14946317 | chr03_14946317 | 7.71 | 0.07 | OsIRO3 | 4 | 27992334 | 27992334 | chr04_27992334 | 12.03 | 3.52 | OsHMA5; OsGPX1; cZOGT1; Kala4; OsPIP1; 2; SMG2; OsMKKK10; OsCPK12; es4 | 5 | 16703102 | 16703102 | chr05_16703102 | 9.5 | 0.11 | YGL1; ygl80; OsEm1; EMP1; OsImp | 6 | 4745474 | 4831914 | chr06_4831914 | 13.67 | 3.29 | OsPTF1; OsERF71; RSUS2; RSs1; SUS2; OsKASI; OsARF16 | 7 | 9578963 | 9578963 | chr07_9578963 | 6.92 | 0.30 | Ghd7 | | 7 | 26527050 | 26562456 | chr07_26527050 | 5.18 | 0.85 | OsLti6a; OsPDK1; Os-eIF6; 1; OsPRMT3; OsHsfB4b | 7 | 26965963 | 26965963 | chr07_26965963 | 6.21 | 0.11 | OsbZIP60; qHMS7-ORF1; qHMS7-ORF3; CYP734A5 | 10 | 2918961 | 2918961 | chr10_2918961 | 7.15 | 2.75 | OsDIL; OsLTP6; OsPRP2; OsPRP1 | 11 | 17832018 | 17832018 | chr11_17832018 | 5.62 | 0.91 | OsHSD1; OsSOT1; STV11 | 12 | 730797 | 730797 | chr12_730797 | 5.64 | 4.08 | OsCIPK14 | 12 | 3864512 | 3864512 | chr12_3864512 | 8.41 | 0.10 | Osmyb2; OsGRX27; OsALDH3H1; OsAPx7; OsAPx6; OsAPx5 | 12 | 11533522 | 11533522 | chr12_11533522 | 8.44 | 0.15 | | SCA | 2 | 2645592 | 2789913 | chr02_2645592 | 5.82 | 0.58 | OseIF4A; OsMAPK33; OsMPK14; OsMPK3; OsPP18; OsRBCS1; LC2; OsVIL3; YL1; LRK1 | 2 | 4255102 | 4310130 | chr02_4255102 | 6.38 | 3.19 | OsDPK2; Os4CL3; OsGL1-2 | 2 | 8481676 | 8481676 | chr02_8481676 | 6.7 | 1.13 | GluD; GluD-1; GluB1; GluB1; RPBF; OsDof3; OsDof-10; OsDof7 | 2 | 29065323 | 29065323 | chr02_29065323 | 7.06 | 2.23 | OSOTP51; OsABA8ox1 | 2 | 35116043 | 35133182 | chr02_35116043 | 7.35 | 0.89 | OsABC1-4; OsNDUFA9; flo13; OsIAA10; RTBP1; CYP97A4; OsCBSX3; OsLpa1 | 3 | 23813743 | 23813743 | chr03_23813743 | 5.75 | 0.04 | | 4 | 18157266 | 18216943 | chr04_18216943 | 5.31 | 0.11 | | 4 | 18903071 | 18907437 | chr04_18903071 | 6.46 | 2.19 | OsExo70‐F3 | 4 | 19647607 | 19647607 | chr04_19647607 | 6.36 | 0.30 | OsAFB2; OsJAZ5; OsTIFY9; LAX2; Gnp4; OsClpC1; OsRap2.6; pi21 | 4 | 19894988 | 20618276 | chr04_19931704 | 8.65 | 0.66 | pi21; OsHAK1; OsClpD2; OSINV4; GIF1; OsCIN2; ACL1; COPT4 | 4 | 20879608 | 21022879 | chr04_20890026 | 7.76 | 1.05 | Asr4; ASR6; OsASR2 | 4 | 22158990 | 22189682 | chr04_22164037 | 5.6 | 0.33 | OsRR1; OsFRO1; OsAP37; OSZEP1 | 4 | 23386304 | 23437449 | chr04_23386304 | 5.86 | 2.07 | MOF; OsGASR2; d11; CYP724B1; CPB1; sg4; cl4; OsMYB80; OsMYB103 | 4 | 23998714 | 23998714 | chr04_23998714 | 7.28 | 1.17 | Rf17; RMS; CRC1; OsPIMT2; OsWS1; WSL3 | 4 | 24214858 | 24214858 | chr04_24214858 | 5.03 | 0.03 | OsPIMT2; OsWS1; WSL3; OsRAD21-2 | | 4 | 27009473 | 27236519 | chr04_27058044 | 9.28 | 0.10 | Glup3; OsVPE1; OsALDH3B1; OsLCBK2; Oshox22; OsYSL16; SDG703; OsERF1 | 4 | 35115325 | 35115325 | chr04_35115325 | 6.27 | 2.23 | OsBSK3; OsCAF1B | 5 | 2264054 | 2450291 | chr05_2284407 | 8.17 | 0.68 | OsSAMS1; OsPti1a; OsLti6b; ddOs32; APG; OsMYB2P-1; OsMPK14; OsGH3-6; OsPPKL2; PTB1; OsVIL4 | 5 | 3049063 | 3158516 | chr05_3111450 | 6.4 | 1.22 | OsGRX15; OsTIR1; APIP6; SRS3; OsKinesin-13A; sar1; Chalk5 | 5 | 3511793 | 3814100 | chr05_3607357 | 7.21 | 1.99 | Chalk5; GS5; OsZIP6 | 5 | 17826917 | 17826917 | chr05_17826917 | 5.04 | 0.87 | OsBiP3; RH1; OsFTIP7; OsRLCK185 | 5 | 18633931 | 18854859 | chr05_18633931 | 5.66 | 0.90 | OsBC1L4; OsNADK3; SMOS1; shb; RLA1 | 5 | 20101785 | 20464861 | chr05_20101785 | 7.02 | 0.91 | TCD5; OsbZIP39; OsPCS1; TCM5; SERF1; OsCc1 | 5 | 21819212 | 23047221 | chr05_22813721 | 6.34 | 1.09 | OsAUX1; OsCOI1b; SH5; OsP5CS; OsP5CS1; OsALDH18B1; Osrboh4; OsAMT2; 1; OsPDC1; OsZIP5 | 5 | 23534940 | 23549991 | chr05_23545588 | 6.84 | 1.22 | OsAMTR1; OsSTN8; EUI1; i-sd-1(t) | 5 | 23946757 | 23946757 | chr05_23946757 | 5.85 | 3.93 | OsDMI3; Osmyb5; SDG728; SDG728; OsCML9; OsCam2 | 6 | 2160801 | 2160801 | chr06_2160801 | 5.12 | 0.02 | dp1; DDF1; OsENOD93a; Hd17; Ef7; OsELF3; OsELF3-1; OsELF3.1; Hd-q; OsPRMT10; SPDT | 6 | 21268145 | 21268145 | chr06_21268145 | 5.23 | 0.56 | | 6 | 25011887 | 25011887 | chr06_25011887 | 5.46 | 0.86 | OsARID3; OsbZIP50; OsbZIP74; TGW6; SPP | 6 | 25716036 | 25716036 | chr06_25716036 | 5.46 | 0.86 | OsPSK | 6 | 26086130 | 26086130 | chr06_26086130 | 5.35 | 0.52 | DCM1; OsSAL1; OVP1; OsVP1 | 6 | 26870826 | 26894242 | chr06_26884987 | 6.32 | 1.35 | OsRpoTp; OsGL1-3; Os4CL4 | 6 | 27148504 | 27148504 | chr06_27148504 | 6.32 | 0.38 | OsGRX18; OsPIN2; OsPLS1; OsbZIP52; RISBZ5 | 7 | 7918837 | 7918837 | chr07_7918837 | 5.29 | 0.11 | OsAPL4; OsAGPL4 | 7 | 8181872 | 8485133 | chr07_8181872 | 5.29 | 0.76 | OsAPL4; OsAGPL4; LLB; pls3; OsMTS1 | 7 | 9129150 | 9129150 | chr07_9129150 | 5.32 | 0.15 | OsNRAMP1; Ghd7 | | 7 | 10123301 | 10123301 | chr07_10123301 | 5.32 | 0.24 | | 7 | 26337284 | 26337284 | chr07_26337284 | 7.73 | 2.36 | RNP29; OsRH36; OsLti6a; OsPDK1 | 7 | 27649737 | 27670134 | chr07_27649737 | 5.09 | 1.25 | NTRC; SPIN6; Fd-GOGAT1; lc7; ABC1; spl23 | 8 | 6184657 | 6293033 | chr08_6184657 | 5.76 | 0.73 | OsHAK12; SPL29; UAP1; OsZIP4 | 8 | 7502244 | 7502244 | chr08_7502244 | 5.76 | 0.36 | OsVIL1; OsDPK1; zl16; OsHAD1 | 8 | 9334312 | 9354921 | chr08_9354921 | 7.8 | 1.41 | | 8 | 15462160 | 15462160 | chr08_15462160 | 5.66 | 3.11 | OsBRL3; HDA705 | 8 | 20967437 | 20988336 | chr08_20981726 | 5.25 | 0.17 | GF14c; 14-3-3; OsMYB106; ONAC106 | 10 | 1520673 | 1520673 | chr10_1520673 | 6.73 | 0.03 | OsSNDP1; OsADF | 10 | 15824748 | 15824748 | chr10_15824748 | 6.07 | 0.09 | | 10 | 16100848 | 16100848 | chr10_16100848 | 7.25 | 0.20 | OsPht1; 8; OsPT8; OsHCI1 | 10 | 16334199 | 16334199 | chr10_16334199 | 5.53 | 0.24 | Osgrp-2 | 10 | 22283651 | 22283651 | chr10_22283651 | 6.51 | 0.92 | MYBS3; OsHox1; REL2; CycP4; 1; CYCU4; 1; OsSAPK3; REK; OsCBL1; OsHUB2; FRRP1; OsCAO2 | 10 | 23018816 | 23183833 | chr10_23018816 | 6.96 | 0.16 | OsMYC2; OsITPK5; JMJ706; OsSRO1a; OsCSLD1; rth-2; OsDUR3 | 11 | 1762018 | 1762018 | chr11_1762018 | 5.66 | 0.12 | OsCML2; OsCPK25 | 11 | 10572420 | 10572420 | chr11_10572420 | 5.36 | 0.10 | | 11 | 22419901 | 22419901 | chr11_22419901 | 5.66 | 0.52 | BGL11(t) | 11 | 27487132 | 27487132 | chr11_27487132 | 5.02 | 2.32 | OsGPAT3; OsJAMyb | 11 | 28800508 | 28800508 | chr11_28800508 | 5.02 | 1.27 | DHD1; CycD7; 1; WDL1 | 12 | 14124949 | 14125467 | chr12_14124949 | 7.3 | 3.19 | OsNCED2 | 12 | 16519849 | 16519849 | chr12_16519849 | 5.31 | 0.98 | DEC | BPH | 1 | 33223657 | 34531439 | chr01_34042495 | 7.89 | 1.68 | OsKTN80c; Pish; OsGH3.1; LC1; OsPME1; OsERF3; OsAP37; CycB1; 1; OsLRR1; OsCML1; OsCam3; OsCaM61; SDG709; OsGAmyb; MYBGA; OsNAC4; ONAC068 | | 2 | 24241454 | 24241454 | chr02_24241454 | 5.01 | 7.46 | du3; OsCBP20; OsNST1; BC14; BBM2; LP2 | 2 | 28453213 | 28453213 | chr02_28453213 | 5.07 | 5.16 | OsPIE1; 709 | 2 | 28881879 | 28881879 | chr02_28881879 | 5.33 | 5.16 | AOX1c; OsPUT1; OsBRXL1; OsGRF4; GS2; GL2; PT2; LGS1; OSOTP51; OsABA8ox1 | 2 | 29687730 | 29687730 | chr02_29687730 | 5.72 | 1.27 | Os4CL2; AOX1c; OsPUT1; OsBRXL1; OsGRF4; GS2; OSOTP51; OsABA8ox1; ADL1; OsDEK1; OsGAP1; OsFAD2-1; Ghd2; DTH2 | 3 | 4911745 | 4911745 | chr03_4911745 | 5.46 | 5.40 | OsRab6a; OsDSR4; pRINO1; OsINO1-1 | 3 | 13598822 | 13598822 | chr03_13598822 | 5.04 | 5.80 | OsDIS1; VLN2 | 4 | 4804275 | 4804275 | chr04_4804275 | 5.7 | 4.53 | ETR2; OS-ETR2 | 4 | 5307896 | 5307896 | chr04_5307896 | 5.33 | 0.83 | OsCPS4; OsCyc1; CYP99A3; OsMAS; OsDTS2; OsKSL4; KS4; CYP99A2 | 5 | 16703102 | 16703102 | chr05_16703102 | 5.96 | 0.11 | YGL1; ygl80; OsEm1; EMP1; OsImp | 6 | 17983222 | 17983222 | chr06_17983222 | 7.23 | 0.24 | | 11 | 3977659 | 3977659 | chr11_3977659 | 15.42 | 5.24 | | 11 | 10747918 | 10763678 | chr11_10747918 | 5.13 | 1.20 | BBM1 | 12 | 3970723 | 3970723 | chr12_3970723 | 5.44 | 1.91 | Osmyb2; OsGRX27; OsALDH3H1; OsAPx7; OsAPx6; OsAPx5 | 12 | 10735585 | 10735585 | chr12_10735585 | 5.73 | 0.67 | Pita; Pi-4a; MIR | 12 | 11432955 | 11432955 | chr12_11432955 | 5.47 | 0.62 | OsRBCS3; OsRBCS5; OsRBCS4 | 12 | 11767635 | 11767635 | chr12_11767635 | 5.47 | 0.62 | | 12 | 12314314 | 12314314 | chr12_12314314 | 6.04 | 1.27 | | 12 | 13772007 | 13772007 | chr12_13772007 | 5.47 | 0.62 | | 12 | 14602287 | 14602287 | chr12_14602287 | 11.49 | 4.68 | | 12 | 24447076 | 24447076 | chr12_24447076 | 5.11 | 0.37 | CycD5 |
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附表2
籼稻抽穗期的全基因组关联分析位点
正文中引用本图/表的段落
为了选择更好的特征标记关联模型, 使用GLM、MLM、FarmCPU和Blink模型进行关联分析。QQ-plot结果表明: Blink模型在控制假阳性方面明显优于GLM、MLM和FarmCPU (附图2~附图4)。因此, 对于抽穗期的5个数据集, 选用BLINK模型的GWAS结果, 在V、GCA、TC、SCA和BPH数据集中共检测到104个显著关联位点, 涉及到12条染色体(详见附表2)。其中4号染色体上检测到的显著位点最多, 为16个, 其次是12号染色体上检测到14个, 而在1号、2号、3号、5号、6号、7号、8号、9号、10号和11号染色体上检测到的显著位点分别仅有3、11、6、11、11、9、5、1、8和9个。在不同的数据集中, 重叠的显著关联位点有3组(图3中黑色虚竖线标识), 基因组距离小于200 kb的显著关联位点有10组(图3中蓝色虚竖线标识, 附表3), 共包含25个显著关联位点。其中, 7号染色体上的Ghd7附近在TC和SCA中分别检测显著的显著关联位点; 2号染色体Ghd2附近在BPH (<200 kb)、SCA (<700 kb)和TC (<1.3 Mb)中分别检测到显著的显著关联位点(图3)。这些结果说明, 抽穗期GWAS结果的重复性较高, 这些位点可能为新发现的抽穗期相关的热点区域。
由表2可知, 在亲本品种群体中, 不同基因中单倍型的组合对应水稻抽穗期和产量的表型分布范围分别为86.71~94. 3 d和22.52~32.15 g。其中, SDG724 (Hap.A)_Hd17 (Hap.E)_Ghd7 (Hap.A)的单倍型组合对应的水稻单株籽粒产量可达31.97 g, 抽穗期为94.3 d, 该组合中各基因的单倍型(表2)对应于单个克隆基因的的优异单倍型(表1)。结果表明优异单倍型的聚合的亲本具有更高的产量和更长的抽穗期。
本文的其它图/表
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