甘蓝型油菜苗期耐旱性综合评价与耐旱性鉴定指标筛选

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李阳阳^{1, 2,},
李驰^{1, 2},
任俊洋^{1, 2},
李志^{1, 2},
张晋峰^{1, 2},
吕蓉蓉^{1, 2},
张恒^{1, 2},
吴丹^{1, 2},
王芹^{1, 2},
周清元¹,
殷家明¹,
李加纳^{1, 2},
刘列钊^{1, 2,,},
唐章林^{1, 2,,}
1.西南大学农学与生物科技学院重庆 400715
2.西南大学农业科学研究院重庆 400715
基金项目: 国家重点研发计划项目2018YFD0100500
重庆市社会事业与民生保障科技创新主题专项cstc2016shms-ztzx80010

详细信息

作者简介:李阳阳, 主要研究方向为油菜耐旱性机理研究。E-mail: liyangyangswu@163.com

通讯作者:刘列钊, 主要研究方向为油菜耐旱性机理研究, E-mail: liezhao2003@126.com
唐章林, 主要研究方向为油菜耐旱性品种选育, E-mail: tangzhlin@swu.edu.cn

中图分类号:S326;S565.4

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出版历程

收稿日期:2020-10-29
录用日期:2021-02-10
刊出日期:2021-08-01

Comprehensive evaluation and identification trait selection of drought resistance at the seedling stage of Brassica napus L.

LI Yangyang^{1, 2,},
LI Chi^{1, 2},
REN Junyang^{1, 2},
LI Zhi^{1, 2},
ZHANG Jinfeng^{1, 2},
LYU Rongrong^{1, 2},
ZHANG Heng^{1, 2},
WU Dan^{1, 2},
WANG Qin^{1, 2},
ZHOU Qingyuan¹,
YIN Jiaming¹,
LI Jiana^{1, 2},
LIU Liezhao^{1, 2,,},
TANG Zhanglin^{1, 2,,}
1. College of Agronomy and Biotechnology, Southwest University, Chongqing 400715, China
2. Academy of Agricultural Sciences, Southwest University, Chongqing 400715, China
Funds: the National Key R&D Program of China2018YFD0100500
the Science & Technology Innovation Project of Chongqing Social Undertakings & Livelihood Securitycstc2016shms-ztzx80010

More Information

Corresponding author:LIU Liezhao, E-mail: liezhao2003@126.com;TANG Zhanglin, E-mail: tangzhlin@swu.edu.cn

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摘要
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。
关键词:甘蓝型油菜/
耐旱性/
种质筛选/
鉴定指标/
苗期
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

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图1229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。
Figure1.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.

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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。
Figure2.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.

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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
Table1.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
地上部鲜重 Shoot fresh weight (g·plant^–1)	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
地上部干重 Shoot dry weight (g·plant^–1)	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
地下部鲜重 Root fresh weight (g·plant^–1)	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
地下部干重 Root dry weight (g·plant^–1)	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
叶片过氧化物酶活性 Leaf peroxidase activity (U·g^–1)	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
叶片丙二醛含量 Leaf malonaldehyde content (μmol·g^–1)	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
叶片可溶性蛋白含量 Leaf soluble protein content (mg·g^–1)	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
叶片脯氨酸含量 Leaf proline content (mg·g^–1)	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
叶片相对含水量 Leaf relative water content (%)	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
叶片可溶性糖含量 Leaf soluble sugar content (mg·g^–1)	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.

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表2229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
Table2.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
性状 Trait	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.

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表3229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
Table3.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
材料名称 Accession name	来源地 Source	值 Value	排序 Rank	值 Value	排序 Rank	值 Value	排序 Rank	类群 Group
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.

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表4229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
Table4.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
因子 Factor	特征值 Eigen value	累计贡献率 Cumulative contribution (%)	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
1	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

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表5229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
Table5.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 factors	0.78^**
综合关联度 Comprehensive relation degree	0.74^**	0.94^**
“”表示相关性达P < 0.01显著水平。“” shows significant correlation at P < 0.01 probability level.

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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
Table6.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 function	0.21^**	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 factors	0.13^*	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 degree	0.21^**	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.

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表7229份甘蓝型油菜各性状耐旱系数的相关系数
Table7.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 weight	0.053
地上部干重 Shoot dry weight	0.044	0.676^**
地下部干重 Root dry weight	0.011	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 content	0.038	0.170^*	0.102	–0.011	–0.048	0.065
叶片脯氨酸含量 Leaf proline content	0.083	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.

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