植物研究 ›› 2021, Vol. 41 ›› Issue (5): 775-788.doi: 10.7525/j.issn.1673-5102.2021.05.016
李雁瓷1,2,3, 付乃峰2,3, 孙加芝2,3, 肖艳2,3, 曹建国1, 田代科2,3()
收稿日期:
2020-04-19
出版日期:
2021-09-20
发布日期:
2021-07-05
通讯作者:
田代科
E-mail:dktian@sibs.ac.cn
作者简介:
李雁瓷(1995—),女,硕士研究生,主要从事秋海棠分类研究。
基金资助:
Yan-Ci LI1,2,3, Nai-Feng FU2,3, Jia-Zhi SUN2,3, Yan XIAO2,3, Jian-Guo CAO1, Dai-Ke TIAN2,3()
Received:
2020-04-19
Online:
2021-09-20
Published:
2021-07-05
Contact:
Dai-Ke TIAN
E-mail:dktian@sibs.ac.cn
About author:
LI Yan-Ci(1995—),female,master student,engaged in the classification of begonias.
Supported by:
摘要:
《中国植物志》和Flora of China关于秋海棠(Begonia grandis)的种内分类存在争议,根据两者的分类依据在实践中无法操作,显然该种的种内单元分类还存在问题。为了全面理解秋海棠的种内多样性,在此基础上对其种内单元进行合理分类,以39个秋海棠居群为研究材料,对33个表型性状进行多样性及数量分类分析。通过数量性状变异分析、质量性状遗传多样性分析、主成分分析和R形聚类分析从33个性状中筛选出主要性状进行Q型聚类分析。结果显示:秋海棠的表型多样性丰富,数量性状变异系数为0.077~0.441,质量性状多样性指数为0.287~1.483。主成分分析结果表明:33个性状可综合为6个主成分,其累计贡献率达72.28%。R型聚类分析结果表明:除叶形和花形大小指标相关外,各性状相对独立。Q型聚类结果显示:39个秋海棠居群按雄花大小、叶片大小、植株大小和叶色分为四组,同Flora of China将该种分成3个亚种处理不一致。因此,对秋海棠的种内单元的划分今后有必要综合分子生物学方面的证据来讨论才更合理。有趣的是发现日本秋海棠的自然化居群的祖先极可能为浙江天目山居群。
中图分类号:
李雁瓷, 付乃峰, 孙加芝, 肖艳, 曹建国, 田代科. 秋海棠(Begonia grandis)的种内表型多样性[J]. 植物研究, 2021, 41(5): 775-788.
Yan-Ci LI, Nai-Feng FU, Jia-Zhi SUN, Yan XIAO, Jian-Guo CAO, Dai-Ke TIAN. Intraspecific Phenotypic Diversity in Begonia grandis[J]. Bulletin of Botanical Research, 2021, 41(5): 775-788.
表1
试验材料来源及编码
编号 No. | 居群 Population | 采集地点 Location | 编号 No. | 居群 Population | 采集地点 Location |
---|---|---|---|---|---|
1 | LNLY | 辽宁凌源Lingyuan,Liaoning | 21 | HNHS | 湖南衡山Hengshan,Hunan |
2 | BJXS | 北京香山Xiangshan,Beijing | 22 | JXLS | 江西庐山Lushan,Jiangxi |
3 | TJJX | 天津蓟县 Jixian,Tianjing | 23 | FJTN | 福建泰宁Taining,Fujian |
4 | HEBJJ | 河北井陉 Jingxing,Hebei | 24 | FJYT | 福建永泰Yongtai,Fujian |
5 | HEBXL | 河北兴隆 Xinglong,Hebei | 25 | FJWYS | 福建武夷山Wuyi Mountain,Fujian |
6 | SDTS | 山东泰安 Taian,Shandong | 26 | ZJNB | 浙江宁波 Ningbo,Zhejiang |
7 | HENZJ | 河南焦作 Jiaozuo,Henan | 27 | ZJLA | 浙江临安 Linan,Zhejiang |
8 | SXLS | 山西灵石 Lingshi,Shanxi | 28 | AHXN | 安徽休宁 Xiuning,Anhui |
9 | HBSNJ | 湖北神农架Shennongjia,Hubei | 29 | CQJFS | 重庆金佛山Jinfo Mountain,Chongqi |
10 | HBYX | 湖北岳西 Yuexi,Hubei | 30 | SCYA | 四川雅安 Yaan,Sichuan |
11 | HBHA | 湖北红安 Hongan,Hubei | 31 | SCEMS | 四川峨眉山Emei Mountain,Sichuan |
12 | HBSZ | 湖北随州 Suizhou,Hubei | 32 | YNJL | 云南丽江 Lijiang,Yunnan |
13 | HBZG | 湖北秭归 Zigui,Hubei | 33 | YNKM | 云南昆明Kunming,Yunnan |
14 | HBYC | 湖北宜昌 Yichuang,Hubei | 34 | RBFS | 日本富山 Toyama,Japan |
15 | HBYC | 湖北利川 Lichuan,Hubei | 35 | RBSH | 日本神户 Kobe,Japan |
16 | HNYS | 湖南永顺 Yongshun,Hunan | 36 | RBDF | 日本大分 Oita,Japan |
17 | HNZJJ | 湖南张家界Zhangjiajie,Hunan | 37 | RBKCT | 日本开成町Kaisei,Japan |
18 | HNSZ | 湖南桑植 Sangzhi,Hunan | 38 | RBQM | 日本群马 Gunma,Japan |
19 | HNCD | 湖南常德 Changde,Hunan | 39 | RBLC | 日本镰仓 Kamakura,Japan |
20 | HNLS | 湖南龙山 Longshan,Hunan |
表2
性状及编码
编号 No. | 性状 Morphology | 编码类型 Code | 编码详细情况 Coding details |
---|---|---|---|
1 | 株高Stem height(cm) | N | 测量值 Measured value |
2 | 分枝多少Stem branch number | M | 无(1);少(2);较多(3);多(4) None(1);Less(2);More(3);Most(4) |
3 | 叶柄长Petiole length(cm) | N | 测量值 Measured value |
4 | 叶柄粗Petiole diameter(mm) | N | 测量值 Measured value |
5 | 叶长Leaf length(cm) | N | 测量值 Measured value |
6 | 叶宽Leaf width(cm) | N | 测量值 Measured value |
7 | 叶长宽比Leaf length to leaf width | D | 计算值 Calculated value |
8 | 叶宽侧宽Width of leaf wide-side(cm) | N | 测量值 Measured value |
9 | 下延叶长Leaf lobe length(cm) | N | 测量值 Measured value |
10 | 叶窄侧宽Width of leaf narrow-side(cm) | N | 测量值 Measured value |
11 | 叶宽窄侧比Leaf wide-side width to narrow-side width | D | 计算值 Calculated value |
12 | 下延叶长同叶长比Leaf lobe length to leaf length | D | 计算值 Calculated value |
13 | 叶斑纹 Leaf variegation | B | 无斑(1);有白斑(2) Spotless(1);White spot(2) |
14 | 腹脉颜色Adaxial vein color | M | 绿(1);红(2) Green(1);Red(2) |
15 | 背脉颜色Abaxial vein color | M | 绿(1);红(2) Green(1);Red(2) |
16 | 叶背颜色Color of dorsal leaf blade | M | 绿(1);红(2);红绿(3) Green(1);Red(2);Red- green(3) |
17 | 雄花花柄颜色Color of staminate flower pedicel | M | 白(1);粉白(2);浅粉(3);粉(4);深粉(5);粉绿(6);红(7) White(1);Pink-white(2);Light-pink(3);Pink(4);Deep-pink(5);Pink-green(6);Red(7) |
18 | 雄花花柄长Length of staminate flower pedicel(mm) | N | 测量值 Measured value |
19 | 雄花花冠长Length of staminate flower corolla(mm) | N | 测量值 Measured value |
20 | 雄花花冠宽Width of staminate flower corolla(mm) | N | 测量值 Measured value |
21 | 雄花花冠长宽比Length to width of staminate flower corolla | D | 计算值Calculated value |
22 | 雄花外轮被片颜色Color of staminate flower outer tepal | M | 白(1);粉白(2);浅粉(3);粉(4);深粉(5) White(1);Pink-white(2);Light-pink(3);Pink(4);Deep-pink(5) |
23 | 雄花外轮被片长Length of outer tepa of staminate flower(mm) | N | 测量值Measured value |
24 | 雄花外轮被片宽Width of staminate flower outer tepal(mm) | N | 测量值 Measured value |
25 | 雄花外轮被片长宽比Length to width of staminate flower outer tepal | D | 计算值Calculated value |
26 | 雄花内轮被片颜色Color of staminate flower inner tepal | M | 白(1);粉白(2);浅粉(3);粉(4);深粉(5) White(1);Pink-white(2);Ligh- pink(3);Pink(4);Deep-pink(5) |
27 | 雄花内轮被片长Length of staminate flower inner tepal(mm) | N | 测量值 Measured value |
28 | 雄花内轮被片宽Width of staminate flower inner tepal(mm) | N | 测量值 Measured value |
29 | 雄花内轮被片长宽比Length to width of staminate flower inner tepal | B | 计算值Calculated value |
30 | 雄蕊群高Androecium height(mm) | N | 测量值 Measured value |
31 | 雄蕊群直径Androecium diameter(mm) | N | 测量值 Measured value |
32 | 雄蕊柱长度Staminal column length(mm) | N | 测量值 Measured value |
33 | 雄蕊数Stamen number | N | 测量值 Measured value |
表3
秋海棠数量性状变异分析
数量性状 Quantitative trait | 极大值 Max. | 极小值 Min. | 平均值 Mean | 极差 Range | 标准差 s | 变异系数 CV |
---|---|---|---|---|---|---|
株高Stem height(cm) | 75.500 | 11.500 | 34.249 | 64.000 | 13.600 | 0.397 |
叶柄长Petiole length(cm) | 17.667 | 2.567 | 7.533 | 15.100 | 2.700 | 0.358 |
叶柄粗Petiole width(mm) | 6.417 | 1.500 | 3.209 | 4.917 | 1.039 | 0.324 |
叶长Leaf length(cm) | 27.533 | 5.400 | 13.447 | 22.133 | 3.985 | 0.296 |
叶宽Leaf width(cm) | 20.400 | 3.725 | 9.042 | 16.675 | 2.991 | 0.331 |
叶长宽比Leaf length to leaf width | 1.854 | 1.310 | 1.510 | 0.544 | 0.116 | 0.077 |
叶宽侧宽Width of leaf wide-side(cm) | 10.867 | 2.233 | 5.269 | 8.633 | 1.526 | 0.290 |
下延叶长Leaf lobe length(cm) | 6.983 | 0.500 | 3.314 | 6.483 | 1.240 | 0.374 |
叶窄侧宽Width of leaf narrow-side(cm) | 8.900 | 1.667 | 4.056 | 7.233 | 1.331 | 0.328 |
叶宽窄侧比Leaf wide-side width to narrow-side width | 1.636 | 1.047 | 1.319 | 0.590 | 0.104 | 0.079 |
下延叶长同叶长比Leaf length to leaf lobe length(cm) | 0.329 | 0.093 | 0.242 | 0.236 | 0.046 | 0.190 |
雄花花柄长Length of staminate flower pedicel(mm) | 26.333 | 5.000 | 15.764 | 21.333 | 4.546 | 0.288 |
雄花花冠长Length of staminate flower corolla(cm) | 43.000 | 17.000 | 27.995 | 26.000 | 6.058 | 0.216 |
雄花花冠宽Width of staminate flower corolla(cm) | 31.000 | 11.500 | 21.472 | 19.500 | 3.843 | 0.179 |
雄花花冠长宽比Length to width of staminate flower corolla | 1.667 | 1.059 | 1.302 | 0.608 | 0.144 | 0.111 |
雄花外被片长Length of staminate flower outer tepal(mm) | 21.500 | 9.600 | 14.529 | 11.900 | 2.748 | 0.189 |
雄花外被片宽Width of staminate flower outer tepal(mm) | 19.500 | 6.800 | 13.178 | 12.700 | 2.806 | 0.213 |
雄花外被片长宽比Length to width of staminate flower outer tepal | 1.412 | 0.893 | 1.112 | 0.518 | 0.107 | 0.096 |
雄花内被片长Length of staminate flower inner tepal(mm) | 15.500 | 6.600 | 10.941 | 8.900 | 1.780 | 0.163 |
雄花内被片宽Width of staminate flower inner tepal(mm) | 14.750 | 2.833 | 7.155 | 11.917 | 2.433 | 0.340 |
雄花内被片长宽比Length to width of staminate flower inner tepal | 3.456 | 0.839 | 1.704 | 2.617 | 0.507 | 0.298 |
雄蕊群高Androecium height(mm) | 11.000 | 2.000 | 6.221 | 9.000 | 1.992 | 0.320 |
雄蕊群直径Androecium diameter(mm) | 8.000 | 2.929 | 5.508 | 5.071 | 1.156 | 0.210 |
雄蕊柱长度Staminal column length(mm) | 6.000 | 0.000 | 2.657 | 6.000 | 1.171 | 0.441 |
雄蕊数Stamen number | 130.667 | 19.857 | 66.682 | 110.810 | 28.876 | 0.433 |
表4
秋海棠质量性状频率分布及多样性
质量性状 Qualitative trait | 频率分布Frequency distribution(%) | 遗传多样性指数H′ | ||||||
---|---|---|---|---|---|---|---|---|
1 | 2 | 3 | 4 | 5 | 6 | 7 | ||
分枝多少 Stem branch number | 0.000 | 60.000 | 20.000 | 20.000 | 0.950 | |||
叶斑纹 Leaf variegation | 91.667 | 8.333 | 0.287 | |||||
腹脉颜色 Adaxial vein color | 63.333 | 36.667 | 0.657 | |||||
背脉颜色 Dorsal vein color | 51.667 | 48.333 | 0.693 | |||||
叶背颜色Color of abaxial leaf blade | 55.000 | 40.000 | 0.695 | |||||
雄花花柄颜色Color of staminate flower pedicel | 6.667 | 11.667 | 26.667 | 45.000 | 3.333 | 3.333 | 3.333 | 1.483 |
雄花外被片颜色Color of staminate flower outer tepal | 10.000 | 13.333 | 23.333 | 50.000 | 3.333 | 1.298 | ||
雄花内被片颜色Color of staminate flower inner tepal | 20.000 | 23.333 | 26.667 | 30.000 | 1.375 |
表5
主成分的特征值, 贡献率及特征向量
性状Trait | 主成分Principal component | |||||
---|---|---|---|---|---|---|
PC1 | PC2 | PC3 | PC4 | PC5 | PC6 | |
株高Stem length | 0.51 | 0.03 | -0.16 | 0.38 | 0.07 | -0.15 |
分枝多少Stem branch number | -0.20 | -0.10 | -0.27 | -0.33 | 0.26 | 0.34 |
叶柄长Petiole length | 0.47 | 0.48 | 0.10 | -0.25 | -0.11 | 0.37 |
叶柄粗Petiole width | 0.64 | 0.44 | 0.03 | -0.19 | -0.07 | 0.16 |
叶长Leaf length | 0.70 | 0.55 | -0.35 | 0.13 | -0.01 | 0.16 |
叶宽Leaf width | 0.77 | 0.51 | -0.27 | 0.07 | -0.06 | 0.06 |
叶长宽比Leaf length to leaf width | -0.52 | -0.02 | -0.28 | 0.19 | 0.20 | 0.46 |
叶宽侧宽Width of leaf wide-side | 0.76 | 0.56 | -0.25 | 0.01 | 0.03 | 0.03 |
下延叶长Leaf lobe length | 0.68 | 0.57 | -0.28 | -0.04 | 0.06 | 0.16 |
叶窄侧宽Width of leaf narrow-side | 0.75 | 0.53 | -0.33 | -0.09 | -0.11 | -0.02 |
叶宽窄侧比Wide-side width to narrow-side width of ratio | -0.16 | -0.11 | 0.35 | 0.38 | 0.52 | 0.29 |
下延叶长同叶长比Leaf length to leaf lobe length | 0.22 | 0.05 | 0.01 | -0.60 | -0.32 | -0.58 |
叶斑纹Leaf variegation | -0.18 | 0.51 | 0.04 | 0.44 | 0.31 | -0.44 |
腹脉颜色Abdominal vein color | 0.28 | 0.52 | 0.55 | -0.11 | 0.21 | 0.06 |
背脉颜色Dorsal vein color | 0.14 | 0.56 | 0.60 | 0.11 | 0.12 | 0.02 |
叶背颜色Color of leaf blade abaxially | 0.22 | 0.41 | 0.50 | -0.15 | 0.23 | -0.13 |
雄花花柄颜色Color of staminate flower pedicel | 0.45 | -0.12 | 0.41 | 0.10 | -0.09 | 0.06 |
雄花花柄长Length of staminate flower pedicel | 0.28 | -0.06 | 0.08 | -0.17 | 0.48 | 0.00 |
雄花花冠长Length of staminate flower corolla | 0.84 | -0.38 | -0.15 | 0.16 | -0.03 | 0.02 |
雄花花冠宽Width of staminate flower corolla | 0.73 | -0.46 | -0.23 | 0.04 | 0.18 | 0.04 |
雄花花冠长宽比Length to width of staminate flower corolla | 0.39 | 0.01 | 0.11 | 0.25 | -0.35 | 0.02 |
雄花外被片颜色Color of staminate flower pedicel | 0.40 | -0.01 | 0.57 | 0.32 | -0.09 | -0.13 |
雄花外被片长Length of staminate flower outer tepal | 0.84 | -0.34 | -0.12 | 0.21 | -0.09 | 0.01 |
雄花外被片宽Width of staminate flower outer tepal | 0.86 | -0.31 | -0.20 | 0.09 | 0.15 | -0.13 |
雄花外被片长宽比Length to width of staminate flower outer tepal | -0.29 | 0.00 | 0.24 | 0.24 | -0.59 | 0.32 |
雄花内被片颜色Color of staminate flower inner tepal | 0.44 | -0.09 | 0.25 | 0.46 | -0.45 | 0.08 |
雄花内被片长Length of staminate flower inner tepal | 0.72 | -0.48 | -0.23 | 0.07 | 0.14 | 0.00 |
雄花内被片宽Width of staminate flower inner tepal | 0.74 | -0.37 | 0.25 | -0.28 | -0.04 | 0.11 |
雄花内被片长宽比Length to width of staminate flower inner tepal | -0.48 | 0.29 | -0.37 | 0.45 | 0.21 | -0.32 |
雄蕊群高Androecium height | 0.81 | -0.32 | 0.23 | -0.06 | 0.11 | -0.01 |
雄蕊群直径Androecium diameter | 0.80 | -0.30 | 0.09 | 0.08 | 0.12 | -0.02 |
雄蕊柱长度Staminal column length | 0.56 | -0.26 | 0.32 | -0.18 | 0.32 | 0.02 |
雄蕊数Stamen number | 0.85 | -0.09 | 0.02 | 0.04 | -0.01 | -0.18 |
特征值Eigenvalue | 11.29 | 4.26 | 2.85 | 2.05 | 1.89 | 1.51 |
方差贡献率Variance contribution rate | 34.22 | 12.91 | 8.64 | 6.21 | 5.74 | 4.57 |
累计贡献率Cumulative contribution rate | 34.22 | 47.13 | 55.77 | 61.98 | 67.71 | 72.28 |
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