Comprehensive evaluation and identification trait selection of drought resistance at the seedling stage of Brassica napus L.
-
摘要: 我国油菜生产经常遭受干旱胁迫而影响产量和品质,综合评价油菜种质资源的耐旱性,筛选耐旱种质,确定耐旱性鉴定指标,是耐旱新品种培育和耐旱机理研究的基础性工作。本研究利用229份甘蓝型油菜种质资源,在苗期设置干旱胁迫组和正常灌溉(对照)组,测定地上和地下部鲜重和干重及叶片过氧化物酶活性、丙二醛含量、脯氨酸含量、可溶性蛋白含量、可溶性糖含量和相对含水量10个性状,采用耐旱系数、聚类分析、隶属函数、主成分分析和灰色关联度分析等方法对其耐旱性进行综合评价。结果显示,苗期干旱胁迫使甘蓝型油菜幼苗地上和地下部干重和鲜重及叶片相对含水量和可溶性蛋白含量显著降低,使叶片过氧化物酶活性、丙二醛含量、可溶性糖含量和脯氨酸含量显著升高,而地下部干重在正常灌溉组与干旱胁迫组之间差异不显著。229份种质资源划分为8个类群,各类群表现出不同的耐旱特性。RR002、9801C、炎81-2、07037、浙油758和09-P64-1为耐旱材料,11-P30、CY16PXW-35、08-P35、09-P36、甲972和A148为干旱敏感材料。地上部鲜重、叶片脯氨酸含量和可溶性糖含量可作为甘蓝型油菜苗期综合耐旱性快速、简便、准确的鉴定指标。Abstract: The yield and quality of Brassica napus L. are often affected by drought stress in China. To identify drought-tolerant and drought-sensitive germplasms, screen for indices of drought tolerance to reduce cost and improve efficiency, provide a basis for breeding drought-tolerant varieties, and investigate the mechanisms of drought tolerance, 229 B. napus accessions were subjected to two water treatments (drought stress and well watering) at the seedling stage. The shoot fresh weight, shoot dry weight, root fresh weight, root dry weight, leaf peroxidase activity, malonaldehyde content, proline content, soluble protein content, soluble sugar content, and leaf relative water content were measured. The drought resistance index, clustering analysis, subordinative function, principal component analysis, and gray correlation analysis were used to comprehensively evaluate drought tolerance. The correlation coefficients between the drought resistance index of each trait and the average value of the subordinative function, composite value of the principal component factors, and comprehensive relation degree and those among the drought resistance indexes of the ten traits were calculated to determine the drought tolerance index. The results showed significant differences between the drought-stressed and well-watered plants at the seedling stage. During drought stress, the leaves of drought-stressed plants gradually wilted and curled to different degrees, and the leaves were born slowly and became smaller, while the old leaves turned yellow. After 30 days of stress, the drought-stressed plants were shorter, weaker, and had lower biomass than the well-watered plants. The shoot fresh weight, shoot dry weight, root fresh weight, leaf relative water content and soluble protein content were lower, and the leaf peroxidase activity, malonaldehyde content, proline content, and soluble sugar content increased under drought stress. The root dry weight did not differ between drought stress and well watering. The 229 B. napus accessions were divided into eight groups, and the accessions of each group showed different drought tolerance characteristics. The significant correlation between the average value of the subordinative function, composite value of the principal component factors, and the comprehensive relation degree indicated that the comprehensive evaluation was reliable. Therefore, by means of the subordinative function, principal component analysis, and gray correlation analysis, accessions RR002, 9801C, Yan81-2, 07037, Zheyou758, and 09-P64-1 were identified as drought tolerant. Accessions 11-P30, CY16PXW-35, 08-P35, 09-P36, Jia972, and A148 were drought sensitive. The correlations of shoot fresh weight, shoot dry weight, root fresh weight, root dry weight, leaf proline content, soluble sugar content with the average value of the subordinative function, composite value of the principal component factors, and the comprehensive relation degree were significant with high correlation coefficients, as did those among shoot fresh weight, shoot dry weight, root fresh weight, and root dry weight. The shoot fresh weight, leaf proline content and soluble sugar content can serve as rapid, simple, and accurate traits to identify comprehensive drought tolerance at the seedling stage in B. napus.
-
Key words:
- Brassica napus L. /
- Drought tolerance /
- Germplasm screening /
- Identification index /
- Seedling stage
-
图 1 229份甘蓝型油菜的8个类群各性状耐旱系数值的热图和多重比较
对于同一性状, 不同大、小写字母分别表示类型间差异达P < 0.01和P < 0.05显著水平。SFW: 地上部鲜重; SDW: 地上部干重; RFW: 地下部鲜重; RDW: 地下部干重; POD: 叶片过氧化物酶活性; MDA: 叶片丙二醛含量; PROTEIN: 叶片可溶性蛋白含量; PRO: 叶片脯氨酸含量; RWC: 叶片相对含水量; SUG: 叶片可溶性糖含量。
Figure 1. Heatmap and multiple comparisons for drought resistance index values of traits in 8 groups of 229 Brassic napus accessions
For one trait, different capital and lowercase letters mean significant differences among groups at P < 0.01 and P < 0.05 levels, respectively. SFW: shoot fresh weight; SDW: shoot dry weight; RFW: root fresh weight; RDW: root dry weight; POD: leaf peroxidase activity; MDA: leaf malonaldehyde content; PROTEIN: leaf soluble protein content; PRO: leaf proline content; RWC: leaf relative water content; SUG: leaf soluble sugar content.
图 2 甘蓝型油菜耐旱材料和干旱敏感材料各性状耐旱系数值热图
红线以上为耐旱材料, 红线以下为干旱敏感材料。SFW: 地上部鲜重; SDW: 地上部干重; RFW: 地下部鲜重; RDW: 地下部干重; POD: 叶片过氧化物酶活性; MDA: 叶片丙二醛含量; PROTEIN: 叶片可溶性蛋白含量; PRO: 叶片脯氨酸含量; RWC: 叶片相对含水量; SUG: 叶片可溶性糖含量。
Figure 2. Heatmap of drought resistance index values of traits of drought resistant and sensitive accessions of Brassic napus
Accessions above the red line are drought-resistant and those below the red line are drought-sensitive. SFW: shoot fresh weight; SDW: shoot dry weight; RFW: root fresh weight; RDW: root dry weight; POD: leaf peroxidase activity; MDA: leaf malonaldehyde content; PROTEIN: leaf soluble protein content; PRO: leaf proline content; RWC: leaf relative water content; SUG: leaf soluble sugar content.
表 1 干旱处理和正常灌溉处理下229份甘蓝型油菜各性状的变化
Table 1. Trait changes of 229 Brassic napus accessions under drought stress and well watering conditions
性状
Trait处理
Treatment平均值
Average标准差
Standard error最小值
Min最大值
Max变异系数
Coefficient of variation地上部鲜重
Shoot fresh weight (g·plant–1)正常灌溉Well watering 45.109**(A) 11.407 21.247 81.800 0.253 干旱处理Drought stress 8.656**(B) 2.483 4.327 17.990 0.287 地上部干重
Shoot dry weight (g·plant–1)正常灌溉Well watering 3.063**(A) 0.776 1.170 6.237 0.253 干旱处理Drought stress 1.354**(B) 0.392 0.505 3.288 0.289 地下部鲜重
Root fresh weight (g·plant–1)正常灌溉Well watering 1.241**(A) 0.515 0.173 3.094 0.414 干旱处理Drought stress 0.790**(B) 0.272 0.227 1.621 0.344 地下部干重
Root dry weight (g·plant–1)正常灌溉Well watering 0.333**(A) 0.093 0.089 0.691 0.279 干旱处理Drought stress 0.348**(A) 0.077 0.172 0.628 0.221 叶片过氧化物酶活性
Leaf peroxidase activity (U·g–1)正常灌溉Well watering 5.987**(B) 0.994 3.681 9.723 0.166 干旱处理Drought stress 6.907**(A) 1.284 2.682 10.434 0.186 叶片丙二醛含量
Leaf malonaldehyde content (μmol·g–1)正常灌溉Well watering 0.030**(B) 0.011 0.011 0.073 0.371 干旱处理Drought stress 0.057**(A) 0.025 0.022 0.182 0.447 叶片可溶性蛋白含量
Leaf soluble protein content (mg·g–1)正常灌溉Well watering 7.380**(A) 2.507 2.799 16.195 0.340 干旱处理Drought stress 1.987**(B) 0.697 0.708 4.671 0.351 叶片脯氨酸含量
Leaf proline content (mg·g–1)正常灌溉Well watering 1.786**(B) 1.517 0.125 13.237 0.849 干旱处理Drought stress 3.546**(A) 2.017 0.254 13.271 0.569 叶片相对含水量
Leaf relative water content (%)正常灌溉Well watering 94.117**(A) 4.079 70.086 99.965 0.043 干旱处理Drought stress 72.259**(B) 7.509 54.926 96.282 0.104 叶片可溶性糖含量
Leaf soluble sugar content (mg·g–1)正常灌溉Well watering 16.355**(B) 4.774 6.199 37.745 0.292 干旱处理Drought stress 31.153**(A) 5.971 16.401 50.094 0.192 “**”表示不同材料间差异达P < 0.01显著水平; 不同大写字母表示各性状在干旱处理和正常灌溉间差异达P < 0.01显著水平。“**” shows significant difference at P < 0.01 probability level among accessions. Different capital letters show significant differences at P < 0.01 probability level between drought stress and well watering. 表 2 229份甘蓝型油菜聚类为不同类群数时各性状耐旱系数值差异的F值
Table 2. F-values of differences of trait drought resistance indexes when 229 Brassic napus accessions clusting-analyzed into different cluster group numbers
性状
Trait类群数Group number 2 3 4 5 6 7 8 叶片过氧化物酶活性Leaf peroxidase activity 0.489 0.715 12.104** 9.059** 7.548** 6.453** 12.331** 地上部鲜重Shoot fresh weight 26.750** 33.086** 58.725** 44.722** 36.595** 30.552** 40.273** 地上部干重Shoot dry weight 46.248** 65.830** 63.557** 48.365** 38.818** 32.323** 29.134** 地下部干重Root dry weight 38.043** 67.687** 59.565** 44.475** 36.575** 42.757** 40.889** 地下部鲜重Root fresh weight 11.921** 24.513** 19.320** 14.526** 11.577** 19.368** 17.046** 叶片丙二醛含量Leaf malonaldehyde content 50.342** 82.932** 56.235** 60.838** 48.967** 40.951** 35.821** 叶片可溶性蛋白含量Leaf soluble protein content 0.006 0.185 9.628** 7.340** 49.138** 42.310** 36.506** 叶片脯氨酸含量Leaf proline content 17.476** 17.839** 14.067** 117.741** 94.409** 78.327** 67.303** 叶片相对含水量Leaf relative water content 0.228 0.248 0.781 1.117 1.017 1.046 3.697** 叶片可溶性糖含量Leaf soluble sugar content 0.371 0.189 3.303* 2.797* 4.261** 18.019** 15.594** “**”和“*”分别表示各类群间性状差异达P < 0.01和P < 0.05显著水平。“**” and “*” show significant difference in traits among groups at P < 0.01 and P < 0.05 probability levels, respectively. 表 3 229份甘蓝型油菜的部分极端材料的平均隶属函数值、主成分因子综合值、综合关联度和类群
Table 3. Average values of subordinative function, composite values of principal component factors and comprehensive relation degrees of some extreme materials in 229 Brassic napus accessions and their groups
材料名称
Accession name来源地
Source平均隶属函数值
Average value of subordinative function主成分因子综合值
Composite value of principal component factors综合关联度
Comprehensive relation degree类群
Group值
Value排序
Rank值
Value排序
Rank值
Value排序
Rank科里纳Kelina 重庆Chongqing 0.1033 227 0.7811 217 0.8772 222 Ⅱ 炎81-2 Yan81-2 重庆Chongqing 0.3148 8 1.3903 8 0.8983 6 Ⅲ SWU40 重庆Chongqing 0.2961 10 1.2003 32 0.8937 23 Ⅰ SWU44 重庆Chongqing 0.3381 4 1.2326 25 0.8962 13 Ⅰ SWU59 重庆Chongqing 0.2400 37 1.3770 11 0.8948 15 Ⅷ CY12PXW-6 四川Sichuan 0.3500 2 1.2908 21 0.8963 12 Ⅲ CY16PXW-35 四川Sichuan 0.1223 222 0.7263 223 0.8770 225 Ⅱ wx10213 湖南Hunan 0.1434 203 0.7196 224 0.8789 216 Ⅱ 10-804 湖南Hunan 0.1411 208 0.6731 227 0.8768 226 Ⅷ 631 湖南Hunan 0.2492 31 1.6645 3 0.9119 3 Ⅶ 1360 湖南Hunan 0.2083 70 1.7604 2 0.9242 2 Ⅶ 07037 湖北Hubei 0.3154 7 1.3807 10 0.8968 10 Ⅲ RR002 湖北Hubei 0.3744 1 1.5993 4 0.9048 4 Ⅲ 宁油1号Ningyou 1 湖北Hubei 0.3331 5 1.3065 18 0.8968 9 Ⅲ 11-9-700 湖北Hubei 0.3039 9 1.3080 17 0.8956 14 Ⅰ 09-P64-1 湖北Hubei 0.3473 3 2.2955 1 0.9571 1 Ⅶ 11-P74-8父本
Male parent of 11-P74-8湖北Hubei 0.1366 214 0.7644 219 0.8780 219 Ⅱ 甲972 Jia 972 湖北Hubei 0.0840 229 0.6112 229 0.8737 229 Ⅱ 甲预05棚Jiayu 05 Peng 湖北Hubei 0.1187 224 0.8147 212 0.8795 215 Ⅱ 浙油758 Zheyou 758 浙江Zhejiang 0.2878 11 1.4124 7 0.8972 8 Ⅵ 垦C1 Ken C1 陕西Shaanxi 0.1250 220 0.7721 218 0.8778 221 Ⅱ GY284 陕西Shaanxi 0.1258 218 0.7384 222 0.8771 224 Ⅳ A82 江西Jiangxi 0.1501 196 0.7552 220 0.8801 210 Ⅴ A148 瑞典Sweden 0.1144 226 0.6761 226 0.8767 227 Ⅳ 08-P35 湖北Hubei 0.1204 223 0.7055 225 0.8772 223 Ⅱ 09-P36 湖北Hubei 0.0952 228 0.6681 228 0.8759 228 Ⅱ 10-P29 湖北Hubei 0.1233 221 0.8305 206 0.8797 214 Ⅱ 11-P30 湖北Hubei 0.1258 219 0.7526 221 0.8780 220 Ⅱ 12-P01 湖北Hubei 0.1168 225 0.8235 209 0.8788 217 Ⅳ 9801C 甘肃Gansu 0.3315 6 1.4178 6 0.8982 7 Ⅲ SWU41 重庆Chongqing 0.2584 23 1.5867 5 0.9014 5 Ⅵ SWU69 重庆Chongqing 0.2381 40 1.3873 9 0.8964 11 Ⅵ 229份材料详情可扫本文首页OSID码查看。Scan the OSID code in the first page of this article to view the details of 229 accessions. 表 4 229份甘蓝型油菜各主成分因子的特征值、贡献率和特征向量值
Table 4. Eigen values of all indexes and their contributions and loading matrix of principal component factors of 229 Brassic napus accessions
因子
Factor特征值
Eigen value累计贡献率
Cumulative contribution (%)特征向量值Eigen vector 叶片过氧化物酶活性
Leaf peroxidase activity地上部鲜重
Shoot fresh weight地上部干重
Shoot dry weight地下部干重
Root dry weight地下部鲜重
Root fresh weight叶片丙二醛含量
Leaf malonaldehyde content叶片可溶性蛋白含量
Leaf soluble protein content叶片脯氨酸含量
Leaf proline content叶片相对含水量
Leaf relative water content叶片可溶性糖含量
Leaf soluble sugar content1 2.630 26.302 0.026 0.511 0.494 0.544 0.431 –0.004 0.069 0.035 0.075 –0.031 2 1.476 41.059 –0.350 –0.178 –0.196 0.145 0.278 0.391 –0.053 –0.426 0.429 0.434 3 1.151 52.564 –0.024 0.163 0.096 –0.160 –0.266 0.412 0.698 0.363 0.230 0.158 4 1.012 62.680 0.668 –0.075 –0.054 0.052 0.055 –0.324 0.314 –0.286 –0.022 0.503 5 0.913 71.807 0.262 –0.133 –0.003 0.068 0.158 0.379 –0.423 0.568 –0.236 0.428 6 0.878 80.591 0.357 0.077 –0.088 –0.013 –0.118 –0.109 –0.310 0.205 0.813 –0.183 7 0.820 88.792 0.479 –0.040 –0.111 0.011 0.141 0.609 0.057 –0.314 –0.128 –0.498 8 0.596 94.749 –0.058 –0.242 –0.485 0.123 0.582 –0.211 0.340 0.370 0.040 –0.224 9 0.298 97.730 0.018 –0.729 0.644 –0.018 0.097 –0.026 0.093 0.035 0.140 –0.120 10 0.227 100.000 0.016 0.245 0.189 –0.796 0.507 –0.025 –0.062 –0.048 0.049 0.059 表 5 229份甘蓝型油菜的平均隶属函数值、主成分因子综合值和综合关联度的相关性
Table 5. Correlations among average value of subordinative function, composite value of principal component factors and comprehensive relation degree of 229 Brassic napus accessions
性状
Trait平均隶属函数值
Average value of subordinative function主成分因子综合值
Composite value of principal component factors主成分因子综合值
Composite value of principal component factors0.78** 综合关联度
Comprehensive relation degree0.74** 0.94** “**”表示相关性达P < 0.01显著水平。“**” shows significant correlation at P < 0.01 probability level. 表 6 甘蓝型油菜各性状耐旱系数值与平均隶属函数值、主成分因子综合值和综合关联度的相关性
Table 6. Correlations between drought resistance index value of each trait and average value of subordinative function, composite value of principal component factors and comprehensive relation degree
性状
Trait叶片过氧化物酶活性
Leaf peroxidase activity地上部鲜重
Shoot fresh weight地上部干重
Shoot dry weight地下部干重
Root dry weight地下部鲜重
Root fresh weight叶片丙二醛含量
Leaf malonaldehyde content叶片可溶性蛋白含量
Leaf soluble protein content叶片脯氨酸含量
Leaf proline content叶片相对含水量
Leaf relative water content叶片可溶性糖含量
Leaf soluble sugar content平均隶属函数值
Average value of subordinative function0.21** 0.69** 0.66** 0.69** 0.56** 0.27** 0.42** 0.22** 0.25** 0.22** 主成分因子综合值
Composite value of principal component factors0.13* 0.40** 0.42** 0.53** 0.47** 0.18** 0.11 0.62** 0.13 0.42** 综合关联度
Comprehensive relation degree0.21** 0.38** 0.39** 0.44** 0.40** 0.15* 0.15* 0.70** 0.09 0.26** “**”和“*”分别表示相关性达P < 0.01和P < 0.05显著水平。“**” and “*” show significant correlation at P < 0.01 and P < 0.05 levels, respectively. 表 7 229份甘蓝型油菜各性状耐旱系数的相关系数
Table 7. Correlation coefficients of trait drought resistance index values in 229 Brassic napus accessions
性状
Trait叶片过氧化物酶活性
Leaf peroxidase activity地上部鲜重
Shoot fresh weight地上部干重
Shoot dry weight地下部干重
Root dry weight地下部鲜重
Root fresh weight叶片丙二醛含量
Leaf malonaldehyde content叶片可溶性蛋白含量
Leaf soluble protein content叶片脯氨酸含量
Leaf proline content叶片相对含水量
Leaf relative water content地上部鲜重
Shoot fresh weight0.053 地上部干重
Shoot dry weight0.044 0.676** 地下部干重
Root dry weight0.011 0.592** 0.571** 地下部鲜重
Root fresh weight–0.031 0.343** 0.314** 0.691** 叶片丙二醛含量
Leaf malonaldehyde content–0.130* –0.046 –0.049 0.004 0.071 叶片可溶性蛋白含量
Leaf soluble protein content0.038 0.170* 0.102 –0.011 –0.048 0.065 叶片脯氨酸含量
Leaf proline content0.083 0.140* 0.133* –0.057 –0.114 –0.006 0.028 叶片相对含水量
Leaf relative water content–0.090 0.087 –0.033 0.124 0.080 0.133 0.033 –0.093 叶片可溶性糖含量
Leaf soluble sugar content–0.034 –0.151* –0.074 0.043 0.071 0.107 0.059 –0.089 0.119 “**”和“*”分别表示相关性达P < 0.01和P < 0.05显著水平。“**” and “*” show significant correlation at P < 0.01 and P < 0.05 levels, respectively. -
[1] ZHU J K. Abiotic stress signaling and responses in plants[J]. Cell, 2016, 167(2): 313-324 doi: 10.1016/j.cell.2016.08.029 [2] BAC-MOLENAAR J A, GRANIER C, KEURENTJES J J, et al. Genome-wide association mapping of time-dependent growth responses to moderate drought stress in Arabidopsis[J]. Plant, Cell & Environment, 2016, 39(1): 88-102 http://www.ncbi.nlm.nih.gov/pubmed/26138664 [3] MARSHALL A, AALEN R B, AUDENAERT D, et al. Tackling drought stress: RECEPTOR-LIKE KINASES present new approaches[J]. The Plant Cell, 2012, 24(6): 2262-2278 doi: 10.1105/tpc.112.096677 [4] MITTLER R. Oxidative stress, antioxidants and stress tolerance[J]. Trends in Plant Science, 2002, 7(9): 405-410 doi: 10.1016/S1360-1385(02)02312-9 [5] LI L H, YI H L. Effect of sulfur dioxide on ROS production, gene expression and antioxidant enzyme activity in Arabidopsis plants[J]. Plant Physiology and Biochemistry, 2012, 58: 46-53 doi: 10.1016/j.plaphy.2012.06.009 [6] CAO Y, LUO Q X, TIAN Y, et al. Physiological and proteomic analyses of the drought stress response in Amygdalus Mira (Koehne) roots[J]. BMC Plant Biology, 2017, 17(1): 53 doi: 10.1186/s12870-017-1000-z [7] HATAMI M, HADIAN J, GHORBANPOUR M. Mechanisms underlying toxicity and stimulatory role of single-walled carbon nanotubes in Hyoscyamus niger during drought stress simulated by polyethylene glycol[J]. Journal of Hazardous Materials, 2017, 324: 306-320 doi: 10.1016/j.jhazmat.2016.10.064 [8] HE F, SHENG M, TANG M. Effects of Rhizophagus irregularis on photosynthesis and antioxidative enzymatic system in Robinia pseudoacacia L. under drought stress[J]. Frontiers in Plant Science, 2017, 8: 183 http://pubmedcentralcanada.ca/pmcc/articles/PMC5311038/ [9] SHEIKH MOHAMMADI M H, ETEMADI N, ARAB M M, et al. Molecular and physiological responses of Iranian perennial ryegrass as affected by trinexapac ethyl, paclobutrazol and abscisic acid under drought stress[J]. Plant Physiology and Biochemistry, 2017, 111: 129-143 doi: 10.1016/j.plaphy.2016.11.014 [10] 路之娟, 张永清, 张楚, 等. 不同基因型苦荞苗期抗旱性综合评价及指标筛选[J]. 中国农业科学, 2017, 50(17): 3311-3322 doi: 10.3864/j.issn.0578-1752.2017.17.006LU Z J, ZHANG Y Q, ZHANG C, et al. Comprehensive evaluation and indicators of the drought resistance of different genotypes of Fagopyrum tataricum at seedling stage[J]. Scientia Agricultura Sinica, 2017, 50(17): 3311-3322 doi: 10.3864/j.issn.0578-1752.2017.17.006 [11] 王璐璐, 杨斌, 肖华贵, 等. PEG-6000模拟干旱胁迫下油菜的根系特性与抗旱性[J]. 种子, 2017, 36(8): 93-95 doi: 10.3969/j.issn.1005-2690.2017.08.065WANG L L, YANG B, XIAO H G, et al. Root system characteristics and drought resistance of rape under drought stress by PEG-6000[J]. Seed, 2017, 36(8): 93-95 doi: 10.3969/j.issn.1005-2690.2017.08.065 [12] 胡承伟, 张学昆, 邹锡玲, 等. PEG模拟干旱胁迫下甘蓝型油菜的根系特性与抗旱性[J]. 中国油料作物学报, 2013, 35(1): 48-53 https://www.cnki.com.cn/Article/CJFDTOTAL-ZGYW201301009.htmHU C W, ZHANG X K, ZOU X L, et al. Root structure and drought tolerance of rapeseed under PEG imposed drought[J]. Chinese Journal of Oil Crop Sciences, 2013, 35(1): 48-53 https://www.cnki.com.cn/Article/CJFDTOTAL-ZGYW201301009.htm [13] 庞红喜, 赵兰芝. 干旱胁迫下不同油菜新品种萌发期耐旱性的比较[J]. 安徽农业科学, 2016, 44(19): 38-41 doi: 10.3969/j.issn.0517-6611.2016.19.015PANG H X, ZHAO L Z. Comparison in drought resistance of different rapeseed varieties in germination period under drought stress[J]. Journal of Anhui Agricultural Sciences, 2016, 44(19): 38-41 doi: 10.3969/j.issn.0517-6611.2016.19.015 [14] 董小云, 米超, 刘自刚, 等. PEG模拟水分胁迫对白菜型冬油菜幼苗生长及生长特性的影响[J]. 河南农业大学学报, 2018, 52(3): 313-321 https://www.cnki.com.cn/Article/CJFDTOTAL-NNXB201803002.htmDONG X Y, MI C, LIU Z G, et al. Response of winter rapessed seeding growth and physiological characteristics under PEG drought tolerance[J]. Journal of Henan Agricultural University, 2018, 52(3): 313-321 https://www.cnki.com.cn/Article/CJFDTOTAL-NNXB201803002.htm [15] 陈致富, 李勤菲, 张永晶, 等. 白菜型油菜品种萌发期的抗旱性鉴定与筛选[J]. 植物遗传资源学报, 2015, 16(1): 15-22 https://www.cnki.com.cn/Article/CJFDTOTAL-ZWYC201501004.htmCHEN Z F, LI Q F, ZHANG Y J, et al. Identification and screening of resources with tolerance against drought stress in Brassica rapa during germination stage[J]. Journal of Plant Genetic Resources, 2015, 16(1): 15-22 https://www.cnki.com.cn/Article/CJFDTOTAL-ZWYC201501004.htm [16] 许媛君, 张生萍, 马晓岗, 等. 白菜型油菜种质资源抗旱鉴定[J]. 青海大学学报: 自然科学版, 2016, 34(1): 1-8 https://www.cnki.com.cn/Article/CJFDTOTAL-QHXZ201601001.htmXU Y J, ZHANG S P, MA X G, et al. The drought resistance identification of Brassica rape germplasm[J]. Journal of Qinghai University: Natural Science Edition, 2016, 34(1): 1-8 https://www.cnki.com.cn/Article/CJFDTOTAL-QHXZ201601001.htm [17] 原小燕, 符明联, 李根泽, 等. 甘蓝型与芥菜型油菜种间杂交后代DH系抗旱性评价[J]. 中国油料作物学报, 2015, 37(1): 62-71 https://www.cnki.com.cn/Article/CJFDTOTAL-ZGYW201501010.htmYUAN X Y, FU M L, LI G Z, et al. Evaluation of drought resistance of DH lines from hybrid of B. napus and B. juncea[J]. Chinese Journal of Oil Crop Sciences, 2015, 37(1): 62-71 https://www.cnki.com.cn/Article/CJFDTOTAL-ZGYW201501010.htm [18] 朱宗河, 郑文寅, 张学昆. 甘蓝型油菜耐旱相关性状的主成分分析及综合评价[J]. 中国农业科学, 2011, 44(9): 1775-1787 doi: 10.3864/j.issn.0578-1752.2011.09.003ZHU Z H, ZHENG W Y, ZHANG X K. Principal component analysis and comprehensive evaluation on morphological and agronomic traits of drought tolerance in rapeseed (Brassica napus L. )[J]. Scientia Agricultura Sinica, 2011, 44(9): 1775-1787 doi: 10.3864/j.issn.0578-1752.2011.09.003 [19] 涂玉琴, 汤洁, 涂伟凤, 等. 与蔊菜属间杂交产生的甘蓝型油菜新材料的抗旱性综合评价[J]. 西南农业学报, 2016, 29(7): 1506-1513 https://www.cnki.com.cn/Article/CJFDTOTAL-XNYX201607003.htmTU Y Q, TANG J, TU W F, et al. Comprehensive evaluation of drought resistance of novel Brassica napus germplasm derived from intergeneric hybridizations with Rorippa indica[J]. Southwest China Journal of Agricultural Sciences, 2016, 29(7): 1506-1513 https://www.cnki.com.cn/Article/CJFDTOTAL-XNYX201607003.htm [20] 谢小玉, 张霞, 张兵. 油菜苗期抗旱性评价及抗旱相关指标变化分析[J]. 中国农业科学, 2013, 46(3): 476-485 doi: 10.3864/j.issn.0578-1752.2013.03.004XIE X Y, ZHANG X, ZHANG B. Evaluation of drought resistance and analysis of variation of relevant parameters at seedling stage of rapeseed (Brassica napus L. )[J]. Scientia Agricultura Sinica, 2013, 46(3): 476-485 doi: 10.3864/j.issn.0578-1752.2013.03.004 [21] 桂月晶. 油菜抗旱指标筛选及抗旱相关基因的表达分析[D]. 开封: 河南大学, 2011: 23-59GUI Y J. Screening of drought resistance indexes and expression of drought-resistant related genes in Brassica napus L. [D]. Kaifeng: Henan University, 2011: 23-59 [22] 洪双, 李浩, 许鲲, 等. 甘蓝型油菜微核心种质耐旱鉴定与评价指标筛选[J]. 中国油料作物学报, 2018, 40(2): 209-217 https://www.cnki.com.cn/Article/CJFDTOTAL-ZGYW201802006.htmHONG S, LI H, XU K, et al. Identification of a mini-core collection of Brassica napus accessions for drought tolerance and selection of evaluation indices[J]. Chinese Journal of Oil Crop Sciences, 2018, 40(2): 209-217 https://www.cnki.com.cn/Article/CJFDTOTAL-ZGYW201802006.htm [23] 牛远, 李玲芬, 杨修艳, 等. 氯化胆碱和海藻糖对油菜蕾薹期干旱胁迫的缓解效应研究和耐旱指标筛选[J]. 核农学报, 2020, 34(4): 860-869 https://www.cnki.com.cn/Article/CJFDTOTAL-HNXB202004023.htmNIU Y, LI L F, YANG X Y, et al. Drought tolerance effects of choline chloride and trehalose on rapeseed (Brassica napus L. ) at bud stage under drought stress and selection of related indices[J]. Journal of Nuclear Agricultural Sciences, 2020, 34(4): 860-869 https://www.cnki.com.cn/Article/CJFDTOTAL-HNXB202004023.htm [24] 侯林涛. 甘蓝型油菜种子老化处理后发芽指数QTL定位及生理分析[D]. 重庆: 西南大学, 2017: 18-20HOU L T. Germination index QTL mapping and physiological analysis for Brassica napus L. seed after artificial aging treatment[D]. Chongqing: Southwest University, 2017: 18-20 [25] 王丹丹. 甘蓝型油菜遗传图谱构建及苗期耐旱相关性状的QTL定位[D]. 重庆: 西南大学, 2014: 20-21WANG D D. Mapping and QTL analysis of genes to drought tolerance in Brassica napus L. [D]. Chongqing: Southwest University, 2014: 20-21 [26] 唐启义. DPS数据处理系统: 实验设计、统计分析及数据挖掘[M]. 2版. 北京: 科学出版社, 2010TANG Q Y. DPS Data Processing System: Experimental Design, Statistical Analysis and Data Mining (Second Edition)[M]. Beijing: Science Press, 2010 [27] 郭雪松. 油菜种质资源耐旱性的鉴定[D]. 重庆: 西南大学, 2009: 9-13GUO X S. Evaluation on drought tolerance ability of rapeseed resources[D]. Chongqing: Southwest University, 2009: 9-13 [28] 李莉, 杨雷, 杨莉, 等. 应用灰色关联分析法综合评价草莓种质资源[J]. 河北农业科学, 2008, 12(4): 18-19, 21 https://www.cnki.com.cn/Article/CJFDTOTAL-HBKO200804007.htmLI L, YANG L, YANG L, et al. A comprehensive assessment on strawberry germplasm resource by using the grey relevant analysis method[J]. Journal of Hebei Agricultural Sciences, 2008, 12(4): 18-19, 21 https://www.cnki.com.cn/Article/CJFDTOTAL-HBKO200804007.htm [29] 张朋飞, 武军艳, 孙万仓, 等. 干旱胁迫对白菜型冬油菜苗期生理特性的影响[J]. 西北农业学报, 2015, 24(2): 84-90 https://www.cnki.com.cn/Article/CJFDTOTAL-XBNX201502016.htmZHANG P F, WU J Y, SUN W C, et al. Effect of drought stress on physiological character of winter rape (Brassica campestris L. ) at seedling stage[J]. Acta Agriculturae Boreali-Occidentalis Sinica, 2015, 24(2): 84-90 https://www.cnki.com.cn/Article/CJFDTOTAL-XBNX201502016.htm [30] 王卫芳. 油菜苗期抗旱性评价及生理机制研究[D]. 武汉: 华中农业大学, 2018: 13-18WANG W F. Evaluation and physiological mechanism of drought resistance in rape seedling stage[D]. Wuhan: Huazhong Agricultural University, 2018: 13-18 [31] 孔宪旺, 孙明茂. 水稻不同种质苗期耐旱性综合评价[J]. 江西农业学报, 2019, 31(4): 1-7 doi: 10.3969/j.issn.1001-8581.2019.04.001KONG X W, SUN M M. Comprehensive evaluation of drought tolerance of different rice germplasms at seedling stage[J]. Acta Agriculturae Jiangxi, 2019, 31(4): 1-7 doi: 10.3969/j.issn.1001-8581.2019.04.001 [32] 蔡东芳, 张书芬, 何俊平, 等. 甘蓝型油菜抗旱性鉴定研究进展[J]. 中国农学通报, 2017, 33(28): 7-12 doi: 10.11924/j.issn.1000-6850.casb16090004CAI D F, ZHANG S F, HE J P, et al. Drought resistance identification in Brassica napus[J]. Chinese Agricultural Science Bulletin, 2017, 33(28): 7-12 doi: 10.11924/j.issn.1000-6850.casb16090004 [33] 田宏先, 李小玉, 施毅, 等. 晋北区地方油菜品种苗期抗旱性比较[J]. 山西农业科学, 2020, 48(9): 1411-1417 doi: 10.3969/j.issn.1002-2481.2020.09.11TIAN H X, LI X Y, SHI Y, et al. Comparison of drought resistance of Brassica juncea varieties in northern Shanxi[J]. Journal of Shanxi Agricultural Sciences, 2020, 48(9): 1411-1417 doi: 10.3969/j.issn.1002-2481.2020.09.11 [34] 田宏先, 施毅, 王瑞霞. 春油菜光合作用及其相关生理特性对苗期干旱胁迫的响应[J]. 种业导刊, 2020, (4): 7-12 https://www.cnki.com.cn/Article/CJFDTOTAL-MLWZ202004003.htmTIAN H X, SHI Y, WANG R X. Response of photosynthesis and related physiological characteristics of Brassica juncea to drought stress at seedling stage[J]. Journal of Seed Industry Guide, 2020, (4): 7-12 https://www.cnki.com.cn/Article/CJFDTOTAL-MLWZ202004003.htm [35] 田宏先, 王瑞霞, 施毅, 等. 油菜根系对苗期水分胁迫的形态及生理响应[J]. 农业科技通讯, 2020, (8): 148-153 https://www.cnki.com.cn/Article/CJFDTOTAL-KJTX202008051.htmTIAN H X, WANG R X, SHI Y, et al. Morphological and physiological responses of roots to water stress at seedling stage in Brassica juncea[J]. Bulletin of Agricultural Science and Technology, 2020, (8): 148-153 https://www.cnki.com.cn/Article/CJFDTOTAL-KJTX202008051.htm [36] 原小燕, 铁朝良, 符明联, 等. 甘芥种间杂交后代DH系花期抗旱性评价[J]. 干旱地区农业研究, 2017, 35(2): 79-88 https://www.cnki.com.cn/Article/CJFDTOTAL-GHDQ201702015.htmYUAN X Y, TIE C L, FU M L, et al. Drought resistance evaluation of DH lines from interspecific hybrid of B. napus and B. juncea in flowering stage[J]. Agricultural Research in the Arid Areas, 2017, 35(2): 79-88 https://www.cnki.com.cn/Article/CJFDTOTAL-GHDQ201702015.htm [37] 王瑞霞, 李小玉, 田宏先. 晋北区芥菜型油菜抗旱性鉴定及综合抗旱指标筛选[J]. 中国农业科技导报, 2020, 22(11): 42-51 https://www.cnki.com.cn/Article/CJFDTOTAL-NKDB202011006.htmWANG R X, LI X Y, TIAN H X. Drought resistance identification and comprehensive drought resistance index screening of rapeseed (Brassica juncea L. )in north Shanxi[J]. Journal of Agricultural Science and Technology, 2020, 22(11): 42-51 https://www.cnki.com.cn/Article/CJFDTOTAL-NKDB202011006.htm [38] 陈娇, 谢小玉, 张小短, 等. 甘蓝型油菜苗期抗旱性鉴定及综合抗旱指标筛选[J]. 中国油料作物学报, 2019, 41(5): 713-722 https://www.cnki.com.cn/Article/CJFDTOTAL-ZGYW201905009.htmCHEN J, XIE X Y, ZHANG X D, et al. Seedling drought resistance and parameter screening of rapeseed[J]. Chinese Journal of Oil Crop Sciences, 2019, 41(5): 713-722 https://www.cnki.com.cn/Article/CJFDTOTAL-ZGYW201905009.htm