Phenotypic variations in wild populations of Paris polyphylla var. chinensis
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摘要:目的
综合分析华重楼Paris polyphylla var. chinensis (Franch.) Hara野生种群的表型变异规律，为其种质资源保护、药材资源开发利用和繁育改良提供理论支撑。
方法以我国分布区内7个野生华重楼地理种群为研究对象，对其17个表型性状进行系统测量，利用方差分析、变异系数分析、多重比较分析和相关性分析等多种数理统计分析方法，探讨其表型变异规律及其与地理气候因子的关系。
结果除叶片数和花萼片数外，野生华重楼其他15个表型性状在种群间和种群内有显著或极显著的差异；种群内的变异(方差分量占比为52.19%)大于种群间的变异(方差分量占比为26.39%)，表型性状的平均分化系数为34.27%，种群内变异是华重楼表型变异的主要来源；各表型性状的平均变异系数为17.72%，变异幅度为6.83%~29.95%，花梗长、叶面积、花萼面积和株高的变异幅度远大于平均变异系数，叶片数、花萼片数、果球横径、果球纵径、果球横纵比和种子千粒质量的变异幅度远小于平均变异系数，表明华重楼花、叶的数量指标及果球、种子的性状稳定性大于叶片形态性状的稳定性；相关分析表明，大多数表型因子与地理气候因子之间存在显著或极显著的相关性。
结论广东清远地理种群变异最为丰富，可作为种质资源收集保存的优选对象。
Abstract:ObjectiveTo comprehensively analyze the phenotypic variations in wild populations of Paris polyphylla var. chinensis (Franch.) Hara, and provide a theoretical support for the protection of germplasm resources, development and utilization of medicinal materials, and breeding improvement.
MethodThe seven wild populations of P. polyphylla var. chinensis in China were used as materials. Based on the measurement of 17 phenotypic traits, by the analyses of variance, variation coefficient, multiple comparisons and correlation, the phenotypic variation and its relationship with geographical climate factors were discussed.
ResultExcept the numbers of leaf and calyx, the other 15 phenotypic traits of wild P. polyphylla var. chinensis showed significant or extremely significant differences among and within populations. The variation within populations (Percentage of variance component was 52.19%) was greater than that among populations (Percentage of variance component was 26.39%). The mean phenotypic differentiation coefficient of phenotypic traits was 34.27% and the main source of variation was within populations. The average variation coefficient of each phenotypic trait was 17.72% and variation coefficient ranged from 6.83% to 29.95%. The variation ranges of peduncle length, leaf area, calyx area and plant height were higher than the average variation coefficients of these five traits, while those of leaf number, calyx number, fruit transverse diameter, fruit longitudinal diameter, fruit transverse diameter to longitudinal diameter ratio and seed 1000-grain weight were far less than the mean variation coefficients, indicating that the stabilities of flower and leaf quantitative trait indexes, fruit and seed traits were greater than those of leaf shape traits. Correlation analysis showed that most of the 17 phenotypic traits had significant or extremely significant correlations with the geographical and climatic factors.
ConclusionThe wild populations of P. polyphylla var. chinensis in Qingyuan City, Guangdong Province have the most abundant geographical population variations and can be used as the optimal target for germplasm collection and preservation.
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表 1 7个野生华重楼地理种群的地理及主要气候因子情况
Table 1 Sample collection information of Paris polyphylla var. chinensis from different distribution regions and climatic data at the sampling sites
样品采集点
Sample site种群
Population经度
Longitude纬度
Latitude海拔/m
Altitude年均温度/℃
Mean annual temperature无霜期/d
Frost-free period年日照/h
Annual sunshine hour年降水量/mm
Annual precipitation江西新余
Xinyu in JiangxiJXXY 114°37′22″E 27°31′45″N 376 17.7 270 1 655.4 1 594.8 浙江丽水
Lishui in ZhejiangZJLS 119°04′18″E 28°38′28″N 1 103 11.3 240 1 635.1 1 598.9 四川雅安
Yaan in SichuanSCYA 103°01′57″E 30°30′24″N 1 450 17.0 295 1 519.2 1 350.0 广东清远
Qingyuan in GuangdongGDQY 112°20′11″E 25°10′10″N 648 18.0 210 2 290.0 1 329.0 福建三明
Sanming in FujianFJSM 117°29′18″E 26°33′24″N 548 18.2 260 1 727.1 1 688.0 湖南娄底
Loudi in HunanHNLD 112°02′62″E 27°48′05″N 290 17.1 270 1 726.0 1 455.6 湖北宜昌
Yichang in HubeiHBYC 111°21′19″E 30°27′44″N 400 16.9 275 1 710.5 1 215.5 
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表 2 野生华重楼种群间和种群内全株、果、叶及种子表型性状的方差分析结果1)
Table 2 Variance analysis of phenptypic traits of wholeplant, leaf, fruit and seeds among and within Paris polyphylla var. chinensis
性状
Trait均方 Mean square 随机误差
Random
errorF 种群间
Among
populations种群内
Within
populations种群间
Among
populations种群内
Within
populations株高 Height 1 060.94 226.26 266.13 4.69** 5.97** 地径 Diameter 6.39 1.32 1.62 4.83** 7.13** 叶片数 No. of leaf 0.53 0.78 0.08 0.68 1.07 叶长 Leaf length 5 728.49 601.72 264.04 9.52** 7.18** 叶宽 Leaf width 1 043.13 43.62 51.48 23.91** 20.99** 叶面积 Leaf area 2 831.39 142.52 138.48 19.87** 3.56* 叶片长宽比 Blade aspect ratio 3.73 0.84 0.15 4.45** 5.39** 花萼片数 No. of calyx 4.52 0.55 1.22 0.82 0.79 花萼长 Calyx length 762.13 49.75 46.97 15.32** 10.32** 花萼宽 Calyx width 119.22 9.90 6.86 12.04** 11.31** 花萼面积 Calyx area 46.00 3.32 2.81 13.86** 7.15** 花萼长宽比 Calyx aspect ratio 0.55 0.13 0.03 4.12** 5.01** 花梗长 Pedicel length 152.91 70.67 25.22 3.16* 6.25** 果球横径 Fruit transverse diameter 89.74 18.79 5.95 4.78** 6.17* 果球纵径 Fruit longitudinal diameter 256.18 25.35 19.35 10.10** 8.33** 果球横纵比 Fruit transverse diameter
to longitudinal diameter ratio0.24 0.11 0.02 21.60** 17.05** 种子千粒质量 Seed 1 000 -grain weight 843.51 69.95 229.20 12.06** 10.87** 1)“*”和“**”分别表示在0.05和0.01水平差异显著(单因素方差分析)
1)“*” and “**” mean significant differences at 0.05 and 0.01 levels respectively(One-way ANOVA)
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表 3 野生华重楼7个地理种群20个表型性状的平均值、标准差及多重比较1)
Table 3 Mean, standard deviation and multiple comparison of 20 phenotypic traits in seven geographical populations of Paris polyphylla var. chinensis
种群2)
Population株高/cm
Height地径/mm
Diameter叶片数
No. of leaf叶长/mm
Leaf length叶宽/mm
Leaf width叶面积/cm2
Leaf areaJXXY 53.67±17.64ab 8.22±1.17ab 6.33±0.58a 134.91±21.64ab 44.89±10.01b 48.84±17.70b ZJLS 66.43±8.46a 7.93±0.70abc 6.67±1.15a 121.90±13.43bc 48.31±4.10ab 46.48±7.92b SCYA 69.20±12.51a 8.63±1.33a 7.00±1.00a 148.38±12.29a 51.85±4.90a 60.28±5.99a GDQY 49.63±24.01abc 6.47±0.84bcd 6.67±1.53a 114.58±55.20c 32.73±7.84d 30.50±21.37cd FJSM 35.33±5.81bc 5.29±0.78d 6.33±0.58a 123.02±10.82bc 35.80±3.96cd 34.73±5.96c HNLD 22.13±2.25c 6.05±0.84cd 6.67±0.58a 112.69±10.17c 38.91±5.91c 34.57±6.84c HBYC 25.83±2.25c 5.02±0.59d 5.67±0.58a 93.57±6.93d 35.70±7.53cd 26.35±6.28d 种群2)
Population叶片长宽比
Blade aspect ratio花萼片数
No. of calyx花萼长/mm
Calyx length花萼宽/mm
Calyx width花萼面积/cm2
Calyx area花萼长宽比
Calyx aspect ratioJXXY 3.06±0.37bc 5.67±0.58a 49.91±7.87a 19.13±4.13a 7.68±2.56a 2.66±0.34ab ZJLS 2.53±0.24c 6.67±1.15a 49.34±5.34a 18.46±1.74ab 7.20±1.29a 2.68±0.26a SCYA 2.89±0.41b 5.80±0.84a 52.06±8.90a 19.23±3.43a 8.03±2.56a 2.73±0.35a
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续表 3 Continued table 3 种群2)
Population叶片长宽比
Blade aspect ratio花萼片数
No. of calyx花萼长/mm
Calyx length花萼宽/mm
Calyx width花萼面积/cm2
Calyx area花萼长宽比
Calyx aspect ratioGDQY 3.79±2.18a 5.33±0.58a 38.80±8.31b 13.79±1.90c 4.29±1.37b 2.81±0.42a FJSM 3.46±0.38ab 4.67±0.58a 36.71±8.41b 13.42±2.52c 3.97±1.59b 2.76±0.52a HNLD 2.95±0.45bc 5.67±0.58a 39.23±5.24b 16.56±2.48b 5.18±1.42b 2.39±0.25bc HBYC 2.72±0.51c 4.33±0.58a 34.19±2.86b 13.04±4.75c 4.26±1.16b 2.26±0.48c 种群2)
Population花梗长/cm
Pedicel length果球横径/mm
Fruit transverse diameter果球纵径/mm
Fruit longitudinal diameter果球横纵比
Fruit transverse diameter to longitudinal diameter ratio种子千粒质量/g
Seed 1 000-grain weightJXXY 18.17±5.03ab 30.45±5.25bc 35.93±7.28ab 0.86±0.10d 100.16±3.00a ZJLS 19.60±2.15ab 34.09±2.84ab 39.94±3.37a 0.85±0.04d 72.25±2.55b SCYA 18.60±3.47ab 29.50±4.41c 34.40±4.34bc 0.86±0.08d 73.25±2.55b GDQY 23.00±15.86a 29.79±3.46bc 26.33±4.40d 1.14±0.12b 77.49±10.10b FJSM 7.83±0.91ab 28.92±2.65c 30.47±3.37cd 0.96±0.88cd 95.76±4.17a HNLD 10.43±4.29ab 36.83±6.65a 36.06±5.95ab 1.02±0.11c 109.99±17.00a HBYC 5.80±1.06b 32.68±2.79abc 26.64±3.94d 1.25±0.19a 68.14±3.37b 1)同列数据后的不同小写字母表示差异显著(P<0.05，Duncan’s法)；2)JXXY：江西新余；ZJLS：浙江丽水；SCYA：四川雅安；GDQY：广东清远；FJSM：福建三明；HNLD：湖南娄底；HBYC：湖北宜昌
1) Different lowercase letters in the same column indicated significant differences among different geographical populations (P<0.05, Duncan’s method)；2) JXXY: Xinyu in Jiangxi; ZJLS: Lishui in Zhejiang; SCYA: Ya’an in Sichuan; GDQY: Qingyuan in Guangdong; FJSM: Sanming in Fujian; HNLD: Loudi in Hunan; HBYC: Yichang in Hubei
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表 4 野生华重楼表型性状的方差分量及种群间表型分化系数
Table 4 Variance component and differentiation coefficients of phenotypic traits among and within populations in Paris polyphylla var. chinensis
性状
Trait方差分量
Variance component方差分量占比/%
Percentage of variance component表型分化
系数/%
Phenotypic differentiation
coefficient种群间
Among populations种群内
Within populations随机误差
Random error种群间
Among populations种群内
Within populations随机误差
Random error株高 Height 352.96 464.74 266.13 32.57 42.88 24.55 43.16 地径 Diameter 2.13 2.77 1.62 32.67 42.48 24.85 43.47 叶片数 No. of leaf 0.18 0.72 0.08 18.37 73.47 8.16 20.00 叶长 Leaf length 301.64 829.57 264.04 21.62 59.46 18.92 26.67 叶宽 Leaf width 52.36 88.04 51.48 27.29 45.88 26.83 37.29 叶面积 Leaf area 144.13 262.02 138.48 26.46 48.11 25.43 35.49 叶片长宽比 Blade aspect ratio 0.19 0.97 0.15 14.50 74.05 11.45 16.38 花萼片数 No. of calyx 1.47 1.63 1.22 34.03 37.73 28.24 47.42 花萼长 Calyx length 53.20 90.08 46.96 27.96 47.35 24.68 37.13 花萼宽 Calyx width 7.75 15.81 6.86 25.48 51.97 22.55 32.89 花萼面积 Calyx area 3.14 5.74 2.81 26.86 49.10 24.04 35.36 花萼长宽比 Calyx aspect ratio 0.04 0.16 0.03 17.39 69.57 13.04 20.00 花梗长 Pedicel length 44.11 93.10 25.22 27.16 57.32 15.53 32.15 果球横径 Fruit transverse diameter 8.46 23.80 5.95 22.14 62.29 15.57 26.22 果球纵径 Fruit longitudinal diameter 26.53 41.64 19.35 30.31 47.58 22.11 38.92 果球横纵比 Fruit transverse diameter to longitudinal diameter ratio 0.02 0.03 0.02 28.57 42.86 28.57 40.00 种子千粒质量 Seed 1 000-grain weight 272.61 271.75 229.20 35.24 35.13 29.63 50.08 平均值 Mean 26.39 52.19 21.42 34.27
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表 5 野生华重楼种群17个表型性状变异系数
Table 5 Variation coefficients of 17 phenotypic traits in Paris polyphylla var. chinensis populations
% 性状
Trait种群1) Population 平均值
MeanJXXY ZJLS SCYA GDQY FJSM HNLD HBYC 株高 Height 32.86 12.73 18.08 48.40 16.44 72.02 8.73 29.89 地径 Diameter 14.22 8.86 15.42 27.17 14.79 13.89 11.80 15.16 叶片数 No. of leaf 9.12 17.32 14.29 17.28 9.12 9.12 10.19 12.35 叶长 Leaf length 16.04 11.02 8.28 48.18 8.79 9.03 7.40 15.53 叶宽 Leaf width 22.31 8.48 9.46 23.96 11.05 15.18 21.10 15.93 叶面积 Leaf area 36.24 17.04 9.93 70.08 17.15 19.78 23.81 27.72 叶片长宽比 Blade aspect ratio 12.06 9.35 14.31 57.54 10.96 15.21 18.87 19.76 花萼片数 No. of calyx 10.19 17.32 10.83 14.43 15.75 10.19 17.32 13.72 花萼长 Calyx length 15.76 10.81 17.10 21.42 22.90 13.35 8.38 15.67 花萼宽 Calyx width 21.60 9.42 17.83 13.77 18.75 15.00 36.47 18.98 花萼面积 Calyx area 33.36 17.87 31.85 31.98 39.97 27.35 27.26 29.95 花萼长宽比 Calyx aspect ratio 12.86 9.70 12.68 15.02 19.14 10.61 21.25 14.47 花梗长 Pedicel length 27.71 10.98 18.63 68.72 11.58 41.07 18.25 28.13
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续表 5 Continued table 5 性状
Trait种群1) Population 平均值
MeanJXXY ZJLS SCYA GDQY FJSM HNLD HBYC 果球横径 Fruit diameter 17.26 8.34 14.96 11.63 9.16 18.07 8.55 12.57 果球纵径 Fruit longitudinal diameter 20.25 8.45 12.62 16.72 11.05 16.49 14.79 14.34 果球横纵比 Fruit aspect ratio 11.71 4.73 9.25 10.52 9.26 10.85 15.01 10.19 种子千粒质量 Seed 1 000-grain weight 3.00 3.52 3.52 13.03 4.35 15.45 4.94 6.83 平均值 Mean 18.62 10.94 14.06 29.99 14.72 19.57 16.12 17.72 1) JXXY：江西新余；ZJLS：浙江丽水；SCYA：四川雅安；GDQY：广东清远；FJSM：福建三明；HNLD：湖南娄底；HBYC：湖北宜昌
1) JXXY: Xinyu in Jiangxi；ZJLS: Lishui in Zhejiang；SCYA: Ya’an in Sichuan；GDQY: Qingyuan in Guangdong；FJSM: Sanming in Fujian；HNLD: Loudi in Hunan；HBYC: Yichang in Hubei
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表 6 野生华重楼17个表型性状与生态地理因子的相关系数1)
Table 6 Correlation coefficients between 17 phenotypic traits of Paris polyphylla var. chinensis and ecological geography factors
性状
Trait经度
Longitude纬度
Latitude海拔
Altitude年均温度
Mean annual temperature无霜期
Frost-free period年日照时数
Annual sunshine
hour年降水量
Annual precipitation株高 Height −0.123 0.085 0.671** −0.353 −0.072 −0.109 0.135 地径 Diameter −0.213 0.172 0.530* −0.293 0.135 −0.267 0.157 叶片数 No. of leaf −0.134 −0.074 0.286 −0.077 −0.041 0.007 0.088 叶长 Leaf length −0.206* 0.072 0.354** 0.010 0.200* −0.288** 0.188* 叶宽 Leaf width −0.222** 0.444** 0.530** −0.365** 0.373** 0.535** 0.133 叶面积 Leaf area −0.277** 0.301** 0.512** −0.179* 0.333** −0.425** 0.154 叶片长宽比 Blade aspect ratio 0.010 −0.340** −0.114 0.284** −0.219* 0.324** 0.008 花萼片数 No. of calyx 0.078 −0.230 0.299 −0.317 −0.344 0.152 0.151 花萼长 Calyx length −0.108 0.267** 0.449** −0.291** 0.199* −0.372** 0.210* 花萼宽 Calyx width −0.075 0.199* 0.314** 0.256** 0.226* −0.385** 0.255** 花萼面积 Calyx area −0.136 0.343** 0.386** −0.268** 0.287** −0.435** 0.170 花萼长宽比 Calyx aspect ratio 0.022 −0.186 0.229* −0.011 −0.176 0.118 0.146 花梗长 Pedicel length −0.071 −0.274 0.312 −0.114 −0.377 0.311 −0.036 果球横径 Fruit transverse diameter 0.059 0.081 −0.176 −0.230* 0.037 −0.063 −0.053 果球纵径 Fruit longitudinal diameter −0.027 0.328** 0.115 −0.362** 0.390** −0.495** 0.266* 果球横纵比 Fruit transverse diameter to longitudinal diameter ratio 0.098 −0.337** −0.317** 0.210 −0.490** 0.594** −0.423** 种子千粒质量 Seed 1 000-grain weight 0.232 −0.272 −0.535** 0.306 0.178 −0.135 0.498* 1)“*”和“**”分别表示在0.05和0.01水平(双侧)上显著相关
1)“*”and“**”indicated significant correlations at 0.05 and 0.01 levels (double-tail), respectively
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