中国水稻科学 ›› 2021, Vol. 35 ›› Issue (6): 573-585.DOI: 10.16819/j.1001-7216.2021.210110
李杰1, 田蓉蓉1, 白天亮1, 朱春艳1, 宋佳伟1, 田蕾1,*(), 马帅国1,2, 吕建东1, 胡慧1, 王震宇1, 罗成科1, 张银霞1, 李培富1
收稿日期:
2021-01-12
修回日期:
2021-03-20
出版日期:
2021-11-10
发布日期:
2021-11-10
通讯作者:
田蕾
基金资助:
Jie LI1, Rongrong TIAN1, Tianliang BAI1, Chunyan ZHU1, Jiawei SONG1, Lei TIAN1,*(), Shuaiguo MA1,2, Jiandong LÜ1, Hui HU1, Zhenyu WANG1, Chengke LUO1, Yinxia ZHANG1, Peifu LI1
Received:
2021-01-12
Revised:
2021-03-20
Online:
2021-11-10
Published:
2021-11-10
Contact:
Lei TIAN
摘要:
【目的】通过对水稻剑叶性状的综合评价,明确剑叶相关性状间及与6个农艺性状的关系。检测剑叶相关性状的QTL,为优良株型品种选育,剑叶性状基因的精细定位和克隆奠定基础。【方法】以日本优质粳稻品种越光和葡萄牙粳稻地方种Bertone构建的回交群体两个世代为实验材料,利用BC3F1群体基因型构建遗传连锁图谱;测定亲本和BC3F2群体各株系剑叶SPAD、剑叶长、剑叶宽,计算剑叶长宽比、剑叶面积;利用隶属函数和标准差系数赋予权重法获得剑叶性状综合评价值(D值),分析其与6个农艺性状间的关系。分别利用单标记分析(SPA)和区间作图(IM)检测水稻剑叶相关性状QTL。【结果】在抽穗灌浆期,两亲本剑叶SPAD值呈现先升高后降低的动态变化。BC3F2群体的5个剑叶相关性状变异丰富,总体表现趋向轮回亲本越光。4个剑叶形态性状间相关性均达到极显著水平,与剑叶SPAD的相关性不显著。主成分和逐步线性回归分析表明剑叶宽、剑叶SPAD、剑叶长、剑叶面积是影响剑叶综合评价值(D值)的主要因子。高D值株系的株高、穗长、茎基粗和单株产量均极显著高于低D值株系,两者的分蘖数和有效穗数差异不显著。共检测到18个控制剑叶性状的QTL,分布在水稻第1、4、7和8染色体上,贡献率分布范围为4.00%~28.00%(SPA)和3.41%~27.00%(IM),除qFLSPAD1之外的17个QTL增效基因均来自Bertone。在第8染色体上的RM22720-RM404区间发现1个QTL簇,含6个主效QTL,分别为qFLL8.1、qFLL8.2、qFLA8.1、qFLA8.2、qD8.1和qD8.2。【结论】获得了剑叶宽、剑叶SPAD、剑叶长和剑叶面积4个评价剑叶性状的关键指标;明确了剑叶性状与单株产量之间的正相关关系;检测到18个剑叶相关性状QTL,位于第8染色体RM22720-RM404区间的QTL簇,是影响剑叶性状的1个重要染色体区域。
李杰, 田蓉蓉, 白天亮, 朱春艳, 宋佳伟, 田蕾, 马帅国, 吕建东, 胡慧, 王震宇, 罗成科, 张银霞, 李培富. 水稻回交群体剑叶性状综合评价及QTL定位[J]. 中国水稻科学, 2021, 35(6): 573-585.
Jie LI, Rongrong TIAN, Tianliang BAI, Chunyan ZHU, Jiawei SONG, Lei TIAN, Shuaiguo MA, Jiandong LÜ, Hui HU, Zhenyu WANG, Chengke LUO, Yinxia ZHANG, Peifu LI. Comprehensive Evaluation and QTL Analysis for Flag Leaf Traits Using a Backcross Population in Rice[J]. Chinese Journal OF Rice Science, 2021, 35(6): 573-585.
图1 亲本及BC3F2群体剑叶性状表现 A–2018年Bertone和越光剑叶SPAD值动态变化;B–亲本和BC3F2代表性株系的剑叶形态学特征。K-越光;1~10表示10个BC3F2代表性株系;竖线代表5 cm。
Fig. 1. Flag leaf traits of parents and BC3F2 population. A, Dynamic changes in SPAD values of flag leaf of Bertone and Koshihikari in 2018; B, Morphological traits of flag leaf of parents and representative lines of BC3F2. K, Koshihikari; 1-10, Representative lines of BC3F2; Vertical lines represent 5 cm.
性状 Trait | 亲本 Parent | 世代 Generation | 群体参数 Parameter of two populations | ||||||
---|---|---|---|---|---|---|---|---|---|
越光Koshihikari | Bertone | 均值±标准差 Mean±SD | 分布范围 Range | 变异系数 CV/% | 峰度 Kurtosis | 偏度 Skewness | |||
剑叶SPAD FLSPAD | 35.67 | 44.06** | BC3F1 | 41.16±3.60 | 32.40~52.50 | 8.75 | 0.18 | 0.31 | |
BC3F2 | 36.40±2.71 | 30.40~49.00 | 7.31 | 2.49 | 1.07 | ||||
剑叶长 FLL/cm | 29.28 | 35.38* | BC3F1 | 30.16±5.18 | 15.83~44.53 | 17.18 | -0.02 | -0.24 | |
BC3F2 | 27.76±2.94 | 20.80~40.75 | 10.59 | 2.54 | 0.92 | ||||
剑叶宽 FLW/cm | 1.21 | 1.80** | BC3F1 | 1.20±0.14 | 0.80~1.57 | 11.67 | 0.42 | -0.30 | |
BC3F2 | 1.21±0.08 | 1.00~2.00 | 5.00 | 45.09 | 5.13 | ||||
剑叶长宽比 FLWR | 24.20 | 19.66* | BC3F1 | 25.30±4.43 | 13.61~42.10 | 17.51 | 1.24 | 0.60 | |
BC3F2 | 23.07±2.43 | 16.00~33.96 | 10.64 | 1.66 | 0.45 | ||||
剑叶面积FLA/cm2 | 26.57 | 47.76** | BC3F1 | 27.32±6.23 | 9.50~46.27 | 22.80 | 0.11 | -0.31 | |
BC3F2 | 25.15±3.67 | 17.66~54.53 | 12.81 | 17.77 | 2.85 |
表1 越光和Bertone及其两个回交群体5个剑叶性状表现
Table 1 Performance of five flag leaf traits of Koshihikari, Bertone and two backcross populations.
性状 Trait | 亲本 Parent | 世代 Generation | 群体参数 Parameter of two populations | ||||||
---|---|---|---|---|---|---|---|---|---|
越光Koshihikari | Bertone | 均值±标准差 Mean±SD | 分布范围 Range | 变异系数 CV/% | 峰度 Kurtosis | 偏度 Skewness | |||
剑叶SPAD FLSPAD | 35.67 | 44.06** | BC3F1 | 41.16±3.60 | 32.40~52.50 | 8.75 | 0.18 | 0.31 | |
BC3F2 | 36.40±2.71 | 30.40~49.00 | 7.31 | 2.49 | 1.07 | ||||
剑叶长 FLL/cm | 29.28 | 35.38* | BC3F1 | 30.16±5.18 | 15.83~44.53 | 17.18 | -0.02 | -0.24 | |
BC3F2 | 27.76±2.94 | 20.80~40.75 | 10.59 | 2.54 | 0.92 | ||||
剑叶宽 FLW/cm | 1.21 | 1.80** | BC3F1 | 1.20±0.14 | 0.80~1.57 | 11.67 | 0.42 | -0.30 | |
BC3F2 | 1.21±0.08 | 1.00~2.00 | 5.00 | 45.09 | 5.13 | ||||
剑叶长宽比 FLWR | 24.20 | 19.66* | BC3F1 | 25.30±4.43 | 13.61~42.10 | 17.51 | 1.24 | 0.60 | |
BC3F2 | 23.07±2.43 | 16.00~33.96 | 10.64 | 1.66 | 0.45 | ||||
剑叶面积FLA/cm2 | 26.57 | 47.76** | BC3F1 | 27.32±6.23 | 9.50~46.27 | 22.80 | 0.11 | -0.31 | |
BC3F2 | 25.15±3.67 | 17.66~54.53 | 12.81 | 17.77 | 2.85 |
图S1 亲本及BC3F2群体剑叶性状表现a:2018年Bertone和越光剑叶SPAD值动态变化;b:亲本和BC3F2代表性家系的剑叶形态学特征;c~g: 亲本及BC3F2群体剑叶SPAD、剑叶长、剑叶宽、剑叶长宽比和剑叶面积的箱线图; A: Bertone;B:越光;1~10:10个BC3F2代表性家系;竖线代表5 cm;*表示0.05水平差异显著;**表示0.01水平差异显著;下同
Fig. S1 Parents and BC3F2 population flag leaf characteristicsa: Dynamic changes of SPAD values of Bertone and Koshihikari flag leaf in 2018; b:Morphological characteristics of flag leaf of parents and representative lines of BC3F2; c~g: Boxplots of flag leaf SPAD, flag leaf length, flag leaf width, length to width ratio of flag leaf and flag leaf area of parents and BC3F2 populations ; A: Bertone; B: Koshihikari; 1~10: 10 representative lines of BC3F2; Vertical lines represent 5 cm; *Significant at P<0.05; **Significant at P<0.01; The same as below.
图2 BC3F2群体5个剑叶相关性状及综合评价值(D值)的频率分布 K代表越光。K,Koshihikari.
Fig. 2. Distribution of five flag leaf-related traits and comprehensive evaluation (D value) for BC3F2 population.
性状 Trait | 剑叶SPAD FLSPAD | 剑叶长 FLL | 剑叶宽 FLW | 剑叶长宽比 FLWR |
---|---|---|---|---|
剑叶长 Flag leaf length | 0.089 | |||
剑叶宽 Flag leaf width | 0.052 | 0.190** | ||
剑叶长宽比 Length to width ratio of flag leaf | 0.069 | 0.897** | -0.259** | |
剑叶面积 Flag leaf area | 0.094 | 0.913** | 0.569** | 0.641** |
表2 BC3F2群体剑叶性状间相关性分析
Table 2 Correlation analysis on flag leaf traits in BC3F2 population.
性状 Trait | 剑叶SPAD FLSPAD | 剑叶长 FLL | 剑叶宽 FLW | 剑叶长宽比 FLWR |
---|---|---|---|---|
剑叶长 Flag leaf length | 0.089 | |||
剑叶宽 Flag leaf width | 0.052 | 0.190** | ||
剑叶长宽比 Length to width ratio of flag leaf | 0.069 | 0.897** | -0.259** | |
剑叶面积 Flag leaf area | 0.094 | 0.913** | 0.569** | 0.641** |
性状 Trait | CI1 | CI2 | CI3 |
---|---|---|---|
剑叶SPAD Flag leaf SPAD | 0.147 | 0.092 | 0.985 |
剑叶长 Flag leaf length | 0.991 | -0.123 | -0.046 |
剑叶宽 Flag leaf width | 0.306 | 0.948 | -0.082 |
剑叶长宽比 Length to width ratio of flag leaf | 0.838 | -0.545 | -0.004 |
剑叶面积 Flag leaf area | 0.954 | 0.288 | -0.074 |
特征值 Eigenvalue | 2.709 | 1.302 | 0.984 |
贡献率 Contribution / % | 54.171 | 26.043 | 19.683 |
累计贡献率Cumulative contribution / % | 54.171 | 80.214 | 99.897 |
表3 5个剑叶性状3个主成分的载荷值、特征值及贡献率
Table 3 Load value, eigenvalue and contribution rate of three principal components of five flag leaf traits.
性状 Trait | CI1 | CI2 | CI3 |
---|---|---|---|
剑叶SPAD Flag leaf SPAD | 0.147 | 0.092 | 0.985 |
剑叶长 Flag leaf length | 0.991 | -0.123 | -0.046 |
剑叶宽 Flag leaf width | 0.306 | 0.948 | -0.082 |
剑叶长宽比 Length to width ratio of flag leaf | 0.838 | -0.545 | -0.004 |
剑叶面积 Flag leaf area | 0.954 | 0.288 | -0.074 |
特征值 Eigenvalue | 2.709 | 1.302 | 0.984 |
贡献率 Contribution / % | 54.171 | 26.043 | 19.683 |
累计贡献率Cumulative contribution / % | 54.171 | 80.214 | 99.897 |
性状 Trait | 剑叶SPAD FLSPAD | 剑叶长 FLL | 剑叶宽 FLW | 长宽比 FLWR | 剑叶面积 FLA | D值 D value | 株高 PH | 穗长 PL | 分蘖数 TN | 有效穗数 EP | 单株产量 YPP |
---|---|---|---|---|---|---|---|---|---|---|---|
剑叶长FLL | 0.456* | 1.000 | |||||||||
剑叶宽FLW | 0.206 | 0.438* | 1.000 | ||||||||
剑叶长宽比FLWR | 0.422* | 0.889** | -0.015 | 1.000 | |||||||
剑叶面积FLA | 0.417* | 0.930** | 0.735** | 0.663** | 1.000 | ||||||
D值 D value | 0.594** | 0.934** | 0.683** | 0.697** | 0.978** | 1.000 | |||||
株高PH | 0.455* | 0.699** | 0.377* | 0.606** | 0.670** | 0.702** | 1.000 | ||||
穗长PL | 0.449* | 0.776** | 0.350 | 0.697** | 0.726** | 0.748** | 0.653** | 1.000 | |||
分蘖数TN | 0.321 | 0.264 | 0.159 | 0.202 | 0.252 | 0.300 | 0.177 | -0.010 | 1.000 | ||
有效穗数EP | 0.260 | 0.290 | 0.222 | 0.203 | 0.299 | 0.327 | 0.187 | 0.036 | 0.963** | 1.000 | |
单株产YPP | 0.478** | 0.598** | 0.525** | 0.422* | 0.649** | 0.686** | 0.526** | 0.451* | 0.666** | 0.720** | 1.000 |
茎基粗BCT | 0.502** | 0.825** | 0.513** | 0.675** | 0.823** | 0.847** | 0.797** | 0.750** | 0.228 | 0.254 | 0.680** |
表4 高、低D值株系剑叶相关性状与农艺性状的相关系数矩阵
Table 4 Correlation matrix of flag leaf-related traits and agronomic traits in BC3F2 lines with high and low D values.
性状 Trait | 剑叶SPAD FLSPAD | 剑叶长 FLL | 剑叶宽 FLW | 长宽比 FLWR | 剑叶面积 FLA | D值 D value | 株高 PH | 穗长 PL | 分蘖数 TN | 有效穗数 EP | 单株产量 YPP |
---|---|---|---|---|---|---|---|---|---|---|---|
剑叶长FLL | 0.456* | 1.000 | |||||||||
剑叶宽FLW | 0.206 | 0.438* | 1.000 | ||||||||
剑叶长宽比FLWR | 0.422* | 0.889** | -0.015 | 1.000 | |||||||
剑叶面积FLA | 0.417* | 0.930** | 0.735** | 0.663** | 1.000 | ||||||
D值 D value | 0.594** | 0.934** | 0.683** | 0.697** | 0.978** | 1.000 | |||||
株高PH | 0.455* | 0.699** | 0.377* | 0.606** | 0.670** | 0.702** | 1.000 | ||||
穗长PL | 0.449* | 0.776** | 0.350 | 0.697** | 0.726** | 0.748** | 0.653** | 1.000 | |||
分蘖数TN | 0.321 | 0.264 | 0.159 | 0.202 | 0.252 | 0.300 | 0.177 | -0.010 | 1.000 | ||
有效穗数EP | 0.260 | 0.290 | 0.222 | 0.203 | 0.299 | 0.327 | 0.187 | 0.036 | 0.963** | 1.000 | |
单株产YPP | 0.478** | 0.598** | 0.525** | 0.422* | 0.649** | 0.686** | 0.526** | 0.451* | 0.666** | 0.720** | 1.000 |
茎基粗BCT | 0.502** | 0.825** | 0.513** | 0.675** | 0.823** | 0.847** | 0.797** | 0.750** | 0.228 | 0.254 | 0.680** |
图4 以粳稻品种越光为遗传背景的Bertone渗入系全基因组图示基因型蓝色区域,杂合基因型;红色区域,Bertone纯合基因型;灰白色区域,越光纯合基因型。
Fig. 4. Graphical genotypes of the 260 BC3F1 lines of Bertone with Koshihikari genetic background. The blue regions indicate the heterozygous genotype; the red regions indicate the homozygous genotype of Bertone; the gray regions indicate the homozygous genotype of Koshihikari.
图5 BC3F1群体剑叶相关性状QTL在水稻染色体上的位置和分布实心符号表示SPA检测到的QTL;空心符号表示IM检测到的QTL。
Fig. 5. Location and the distribution of QTLs for flag leaf related traits in BC3F1 population on chromosomes in rice. Solid symbol represents the QTL to which SPA is legally assigned;Hollow symbol represents the QTL to which IM is legally assigned.
QTL | 染色体Chromosome | 位置 Position /cM | 标记 Marker | 标记区间 Marker interval | 单标记分析SPA | 区间作图IM | ||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|
LOD | 贡献率PVE/% | 加性效应 Add | LOD | 贡献率PVE/% | 加性效应 Add | 文献 Reference | ||||||
qFLSPAD1 | 1 | 213.8 | RM297 | RM297—RM5310 | 3.91 | 7.00 | 1.51 | 3.16 | 5.57 | 1.21 | [35] | |
qFLSPAD4.1 | 4 | 0.0 | RM16260 | RM16260—RM7585 | 4.17 | 7.00 | -1.39 | 2.81 | 5.00 | -1.18 | ||
qFLSPAD4.2 | 4 | 66.2 | RM7200 | RM7585—RM401 | 9.91 | 17.00 | -2.83 | 4.47 | 7.69 | -1.46 | ||
qFLL8.1 | 8 | 115.0 | RM22720 | RM22720—RM3395 | 11.65 | 19.00 | -3.63 | 11.61 | 19.00 | -3.50 | ||
qFLL8.2 | 8 | 122.3 | RM404 | RM3395—RM404 | 12.26 | 20.00 | -3.44 | 11.29 | 18.00 | -3.15 | ||
qFLW7.1 | 7 | 91.7 | RM5508 | RM182—RM505 | 2.53 | 4.00 | -0.03 | 2.02 | 3.41 | -0.02 | [26] | |
qFLW7.2 | 7 | 101.7 | RM505 | RM5508—RM18 | 4.24 | 7.00 | -0.03 | 2.60 | 4.41 | -0.03 | [42] | |
qFLW8.1 | 8 | 115.0 | RM22720 | RM22720—RM6032 | 9.74 | 16.00 | -0.07 | 9.11 | 14.83 | -0.06 | ||
qFLW8.2 | 8 | 118.8 | RM6032 | RM6032—RM404 | 6.59 | 11.00 | -0.05 | 6.71 | 11.20 | -0.05 | ||
qFLWR8.1 | 8 | 118.8 | RM6032 | RM22720—RM6032 | 4.22 | 7.00 | -1.65 | 3.70 | 6.33 | -1.64 | ||
qFLWR8.2 | 8 | 122.3 | RM404 | RM6032—RM404 | 5.15 | 9.00 | -1.92 | 4.16 | 7.20 | -1.67 | ||
qFLA1 | 1 | 161.1 | RM9 | RM11245—RM488 | 3.74 | 6.00 | -1.60 | 2.38 | 4.17 | -1.30 | ||
qFLA8.1 | 8 | 117.7 | RM3395 | RM22720—RM6032 | 18.26 | 28.00 | -3.96 | 17.78 | 27.00 | -4.43 | ||
qFLA8.2 | 8 | 122.3 | RM404 | RM6032—RM404 | 16.00 | 25.00 | -4.24 | 15.53 | 24.20 | -4.05 | ||
qFLA8.3 | 8 | 140.8 | GR01 | RM404—RM23170 | 2.63 | 5.00 | -1.41 | 4.33 | 7.26 | -1.75 | ||
qD1 | 1 | 161.1 | RM9 | RM11245—RM488 | 4.17 | 7.00 | -0.06 | 2.53 | 4.35 | -0.05 | ||
qD8.1 | 8 | 117.7 | RM3395 | RM22720—RM6032 | 16.70 | 26.00 | -0.14 | 16.77 | 25.50 | -0.16 | ||
qD8.2 | 8 | 122.3 | RM404 | RM6032—RM404 | 14.72 | 23.00 | -0.15 | 14.89 | 23.20 | -0.14 |
表5 两种方法定位到的水稻剑叶相关性状QTL
Table 5 QTL for traits related to flag leaf of rice detected by two methods.
QTL | 染色体Chromosome | 位置 Position /cM | 标记 Marker | 标记区间 Marker interval | 单标记分析SPA | 区间作图IM | ||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|
LOD | 贡献率PVE/% | 加性效应 Add | LOD | 贡献率PVE/% | 加性效应 Add | 文献 Reference | ||||||
qFLSPAD1 | 1 | 213.8 | RM297 | RM297—RM5310 | 3.91 | 7.00 | 1.51 | 3.16 | 5.57 | 1.21 | [35] | |
qFLSPAD4.1 | 4 | 0.0 | RM16260 | RM16260—RM7585 | 4.17 | 7.00 | -1.39 | 2.81 | 5.00 | -1.18 | ||
qFLSPAD4.2 | 4 | 66.2 | RM7200 | RM7585—RM401 | 9.91 | 17.00 | -2.83 | 4.47 | 7.69 | -1.46 | ||
qFLL8.1 | 8 | 115.0 | RM22720 | RM22720—RM3395 | 11.65 | 19.00 | -3.63 | 11.61 | 19.00 | -3.50 | ||
qFLL8.2 | 8 | 122.3 | RM404 | RM3395—RM404 | 12.26 | 20.00 | -3.44 | 11.29 | 18.00 | -3.15 | ||
qFLW7.1 | 7 | 91.7 | RM5508 | RM182—RM505 | 2.53 | 4.00 | -0.03 | 2.02 | 3.41 | -0.02 | [26] | |
qFLW7.2 | 7 | 101.7 | RM505 | RM5508—RM18 | 4.24 | 7.00 | -0.03 | 2.60 | 4.41 | -0.03 | [42] | |
qFLW8.1 | 8 | 115.0 | RM22720 | RM22720—RM6032 | 9.74 | 16.00 | -0.07 | 9.11 | 14.83 | -0.06 | ||
qFLW8.2 | 8 | 118.8 | RM6032 | RM6032—RM404 | 6.59 | 11.00 | -0.05 | 6.71 | 11.20 | -0.05 | ||
qFLWR8.1 | 8 | 118.8 | RM6032 | RM22720—RM6032 | 4.22 | 7.00 | -1.65 | 3.70 | 6.33 | -1.64 | ||
qFLWR8.2 | 8 | 122.3 | RM404 | RM6032—RM404 | 5.15 | 9.00 | -1.92 | 4.16 | 7.20 | -1.67 | ||
qFLA1 | 1 | 161.1 | RM9 | RM11245—RM488 | 3.74 | 6.00 | -1.60 | 2.38 | 4.17 | -1.30 | ||
qFLA8.1 | 8 | 117.7 | RM3395 | RM22720—RM6032 | 18.26 | 28.00 | -3.96 | 17.78 | 27.00 | -4.43 | ||
qFLA8.2 | 8 | 122.3 | RM404 | RM6032—RM404 | 16.00 | 25.00 | -4.24 | 15.53 | 24.20 | -4.05 | ||
qFLA8.3 | 8 | 140.8 | GR01 | RM404—RM23170 | 2.63 | 5.00 | -1.41 | 4.33 | 7.26 | -1.75 | ||
qD1 | 1 | 161.1 | RM9 | RM11245—RM488 | 4.17 | 7.00 | -0.06 | 2.53 | 4.35 | -0.05 | ||
qD8.1 | 8 | 117.7 | RM3395 | RM22720—RM6032 | 16.70 | 26.00 | -0.14 | 16.77 | 25.50 | -0.16 | ||
qD8.2 | 8 | 122.3 | RM404 | RM6032—RM404 | 14.72 | 23.00 | -0.15 | 14.89 | 23.20 | -0.14 |
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