林业科学 ›› 2021, Vol. 57 ›› Issue (8): 56-67.doi: 10.11707/j.1001-7488.20210806
曹德美1,张亚红1,成星奇1,项晓冬1,张磊1,胡建军1,2,*
收稿日期:
2020-09-02
出版日期:
2021-08-25
发布日期:
2021-09-30
通讯作者:
胡建军
基金资助:
Demei Cao1,Yahong Zhang1,Xingqi Cheng1,Xiaodong Xiang1,Lei Zhang1,Jianjun Hu1,2,*
Received:
2020-09-02
Online:
2021-08-25
Published:
2021-09-30
Contact:
Jianjun Hu
摘要:
目的: 揭示青杨不同种群叶片表型性状的遗传多样性和地理变异规律,为进一步解析叶片性状的遗传机制奠定基础,为青杨种质资源收集、保存和品种改良提供参考。方法: 以来自全国主要分布区的34个青杨种群为材料,采集1~2年生枝条在温室扦插繁殖,移植苗圃,第2年春季平茬幼化,同年7月采集茎尖以下第7、8、9片新生叶测定苗期叶片14个表型性状(叶片形态性状、叶色和叶绿素相对含量),并进行方差分析、相关性分析和聚类分析。结果: 青杨的14个叶片表型性状在种群间和种群内均达到极显著差异(P < 0.01),表明青杨叶片表型性状遗传变异丰富;种群间的平均表型分化系数为55.771%,表明种群间变异是主要的变异来源。各表型性状的平均变异系数为20.562%,变异系数最大的是叶柄相对长,最小的是叶绿素SPAD值。各种群叶片表型性状的平均变异系数为12.852%,其中内蒙古坝底(31号)种群变异最丰富,而河北龙泉(11号)种群变异最小,叶片表型性状稳定性最高。青杨叶片表型性状间及其与地理气候因子间大多达到显著或极显著相关,仅叶色与其他叶片性状无明显相关,叶色遗传相对独立。通过聚类分析可将34个种群划分为4个类群,分别为长叶柄大叶类群、中等叶类群、短叶柄窄叶类群和小叶类群。结合类群分布和中国地形特点,中国青杨种群可划分为3个分布区,分别是太行山以东华北平原北缘地区(1区)、太行山和六盘山-秦岭之间的黄土高原地区(2区)和青藏高原周边地区(3区),其中1区以短叶柄窄叶类群为主(57.14%),2区中等叶类群占66.67%,而3区占比最大的为小叶类群(62.5%)。结论: 青杨叶片表型性状遗传变异丰富,河北蓝旗卡伦(17号)、内蒙古半截塔(30号)和内蒙古坝底(31号)种群多样性丰富,是多样性保护和种质资源保存的重点种群。叶片表型与地理气候因子相关性强,青杨叶片呈现以海拔变化为主的地理变异模式。
中图分类号:
曹德美,张亚红,成星奇,项晓冬,张磊,胡建军. 青杨不同种群叶片表型性状的遗传变异[J]. 林业科学, 2021, 57(8): 56-67.
Demei Cao,Yahong Zhang,Xingqi Cheng,Xiaodong Xiang,Lei Zhang,Jianjun Hu. Genetic Variation of Leaf Phenotypic Traits in Different Populations of Populus cathayana[J]. Scientia Silvae Sinicae, 2021, 57(8): 56-67.
表1
青杨种群的地理信息"
编号 Code | 种群 Population | 样木株数 Number of sample trees | 纬度 Latitude (N)/(°) | 经度 Longitude (E)/(°) | 海拔 Altitude (AL)/m | 年均气温 Mean annual temperature (MAT)/℃ | 年降水量 Mean annual rainfall (MAR)/mm | 年均相对湿度 Mean annual humidity (MAH)(%) |
1 | 北京涧沟Jiangou, Beijing | 12 | 40.07 | 116.05 | 895~1 042 | 12.5 | 568 | 54.7 |
2 | 北京小龙门Xiaolongmen, Beijing | 10 | 39.96 | 115.43 | 953~1 379 | 10.2 | 450 | 53.5 |
3 | 陕西蓝田Lantian, Shaanxi | 11 | 34.00 | 109.46 | 874~1 256 | 13.8 | 569 | 71.1 |
4 | 陕西长安Chang’an, Shaanxi | 8 | 33.97 | 109.00 | 826~1 302 | 12.2 | 775 | 69.8 |
5 | 陕西太白Taibai, Shaanxi | 17 | 34.05 | 107.59 | 1 042~1 084 | 8.1 | 713 | 69.8 |
6 | 四川九寨沟Jiuzhaigou, Sichuan | 13 | 33.06 | 103.92 | 1 108~1 608 | 13.1 | 541 | 64.8 |
7 | 四川永和Yonghe, Sichuan | 14 | 33.30 | 104.39 | 1 611~1 704 | 13.1 | 541 | 64.8 |
8 | 甘肃临夏Linxia, Gansu | 7 | 34.34 | 103.33 | 1 980~2 422 | 7.2 | 377 | 63.2 |
9 | 甘肃张掖Zhangye, Gansu | 12 | 38.25 | 100.92 | 2 758~2 782 | 4.1 | 354 | 52.6 |
10 | 河北易县Yixian, Hebei | 19 | 39.38 | 115.06 | 809~936 | 12.5 | 539 | 62.3 |
11 | 河北龙泉Longquan, Hebei | 19 | 38.92 | 113.78 | 742~869 | 2.1 | 673 | 59.8 |
12 | 河北青龙Qinglong, Hebei | 20 | 40.49 | 118.82 | 738~826 | 9.6 | 656 | 59.3 |
13 | 河北涿鹿Zhuolu, Hebei | 20 | 40.08 | 115.09 | 1 145~1 206 | 9.4 | 366 | 53.8 |
14 | 河北小五台Xiaowutai, Hebei | 15 | 39.94 | 114.96 | 1 283~1 419 | 7.6 | 400 | 54.8 |
15 | 河北丰宁Fengning, Hebei | 17 | 41.49 | 116.76 | 1 072~1 169 | 7.2 | 447 | 53.1 |
16 | 河北苏家店Sujiadian, Hebei | 20 | 41.59 | 116.71 | 1 066~1 233 | 7.2 | 447 | 53.1 |
17 | 河北蓝旗卡伦Lanqikalun, Hebei | 6 | 41.68 | 118.12 | 1 163~1 186 | 5.6 | 435 | 54.4 |
18 | 青海湟源Huangyuan, Qinghai | 8 | 36.68 | 101.26 | 2 630~2 635 | 4.3 | 541 | 59.1 |
19 | 青海湟中Huangzhong, Qinghai | 13 | 36.83 | 101.39 | 2 621~2 659 | 4.3 | 541 | 59.1 |
20 | 青海乐都Ledu, Qinghai | 11 | 36.45 | 102.50 | 1 903~1 941 | 8.3 | 338 | 58.2 |
21 | 青海大通Datong, Qinghai | 11 | 37.01 | 101.79 | 2 625~2 829 | 4.1 | 503 | 62.8 |
22 | 山西广灵Guangling, Shanxi | 13 | 39.67 | 114.02 | 1 491~1 684 | 7.0 | 372 | 55.1 |
23 | 山西方山Fangshan, Shanxi | 16 | 37.84 | 111.37 | 1 573~1 681 | 7.7 | 506 | 56.4 |
24 | 山西牛尾Niuwei, Shanxi | 15 | 38.27 | 112.36 | 1 109~1 245 | 8.5 | 428 | 61.3 |
25 | 山西五台Wutai, Shanxi | 17 | 38.76 | 113.62 | 1 064~1 170 | 2.1 | 673 | 59.8 |
26 | 山西七里峪Qiliyu, Shanxi | 18 | 36.60 | 111.96 | 1 290~1 504 | 12.4 | 435 | 58.8 |
27 | 山西右玉Youyu, Shanxi | 18 | 39.94 | 112.57 | 1 399~1 422 | 4.2 | 407 | 58.8 |
28 | 内蒙古宁城Ningcheng, Inner Mongolia | 20 | 41.54 | 118.39 | 1 077~1 184 | 7.3 | 467 | 53.3 |
29 | 内蒙古美林Meilin, Inner Mongolia | 16 | 41.64 | 118.45 | 1 127~1 207 | 7.3 | 467 | 53.3 |
30 | 内蒙古半截塔Banjieta, Inner Mongolia | 6 | 41.83 | 117.56 | 1 080~1 228 | 5.6 | 435 | 54.4 |
31 | 内蒙古坝底Badi, Inner Mongolia | 9 | 42.06 | 116.96 | 1 228~1 299 | 2.4 | 398 | 61.1 |
32 | 内蒙古围场Weichang, Inner Mongolia | 11 | 42.36 | 117.50 | 1 319~1 497 | 5.6 | 435 | 54.4 |
33 | 内蒙古丰镇Fengzhen, Inner Mongolia | 6 | 40.33 | 113.18 | 1 213~1 221 | 5.7 | 379 | 55.4 |
34 | 内蒙古凉城Liangcheng, Inner Mongolia | 8 | 40.43 | 112.28 | 1 278~1 856 | 5.9 | 406 | 53.5 |
表2
青杨叶片表型性状变异"
性状 Traits | 均值 Mean | 极差 Range | 标准差 Standard deviation (SD) | 种群间变异系数 Coefficient of variation among populations(%) | 种群内变异系数 Coefficient of variation within population(%) | 多样性指数 Shannon- Wiener index (H) | 重复力 Repeatability (rep) |
侧脉数Number of lateral veins(LVN) | 19.400 | 18.556 | 3.416 | 17.610 | 9.320 | 2.073 | 0.959 |
叶基角Leaf base angle(LBA)/(°) | 94.331 | 91.719 | 15.658 | 16.599 | 11.826 | 1.950 | 0.918 |
侧脉夹角Lateral vein angle(LVA)/(°) | 49.638 | 37.178 | 5.440 | 10.958 | 7.912 | 2.043 | 0.912 |
叶面积Leaf area(LA)/cm2 | 118.743 | 254.302 | 37.966 | 31.974 | 23.840 | 2.064 | 0.928 |
叶周长Leaf perimeter(LP)/cm | 52.120 | 66.843 | 9.762 | 18.730 | 13.443 | 2.073 | 0.932 |
叶长Leaf length(LL)/cm | 16.363 | 14.606 | 2.535 | 15.492 | 10.887 | 2.075 | 0.927 |
叶宽Leaf width(LW)/cm | 10.271 | 13.742 | 2.094 | 20.384 | 14.662 | 2.052 | 0.936 |
叶柄长Petiole length(PL)/cm | 3.247 | 4.727 | 1.052 | 32.396 | 20.919 | 2.093 | 0.953 |
叶形指数Leaf index(LI) | 1.642 | 1.604 | 0.272 | 16.537 | 10.430 | 2.021 | 0.936 |
叶柄相对长Relative length of petiole(RPL) | 5.843 | 14.529 | 2.221 | 38.006 | 21.108 | 1.848 | 0.944 |
最大叶宽位置Position of leaf width(LWP) | 0.728 | 0.656 | 0.082 | 11.284 | 7.744 | 2.054 | 0.911 |
叶缘因子Leaf margin factor(LMF) | 0.172 | 0.106 | 0.019 | 10.908 | 8.488 | 2.079 | 0.892 |
叶色Leaf color(LC) | 2.219 | 3.000 | 0.803 | 36.175 | 11.521 | 1.054 | 0.985 |
叶绿素SPAD(SPAD) | 42.013 | 26.240 | 4.544 | 10.816 | 7.821 | 2.068 | 0.841 |
均值Mean | — | — | — | 20.562 | 12.852 | 1.968 | 0.927 |
表3
青杨叶片表型性状的方差分析"
性状 Traits | 方差分量Variance component | 表型分化系数 Differentiation coefficients of phenotypic traits (Vst)(%) | F | |||
种群间 Among populations | 种群内 Within population | 随机误差 Random errors | 种群间 Among populations | 种群内 Within population | ||
LVN | 10 275.03 | 5 182.88 | 2 355.831 | 66.47 | 24.21** | 4.98** |
LBA | 162 368.03 | 161 769.87 | 88 419.845 | 50.09 | 12.26** | 4.14** |
LVA | 18 755.22 | 20 110.94 | 25 341.697 | 48.26 | 11.39** | 1.80** |
LA | 1 011 582.20 | 885 361.13 | 621 401.157 | 53.33 | 13.95** | 3.22** |
LP | 68 765.59 | 56 763.12 | 40 559.040 | 54.78 | 14.79** | 3.17** |
LL | 4 431.90 | 3 958.20 | 2 523.670 | 52.82 | 13.67** | 3.55** |
LW | 3 250.31 | 2 536.08 | 1 622.159 | 56.17 | 15.65** | 3.50** |
PL | 943.32 | 542.18 | 421.016 | 63.50 | 21.25** | 2.91** |
LI | 54.91 | 43.20 | 21.194 | 55.97 | 15.52** | 4.61** |
RPL | 3 905.59 | 2 669.19 | 1 802.884 | 59.40 | 17.87** | 3.35** |
LWP | 4.30 | 4.67 | 3.334 | 47.96 | 11.25** | 3.17** |
LMF | 0.20 | 0.26 | 0.284 | 43.14 | 9.27** | 2.10** |
LC | 707.94 | 132.35 | 2.000 | 84.25 | 65.32** | 149.76** |
SPAD | 8 255.08 | 10 233.59 | 6 214.112 | 44.65 | 6.28** | 3.97** |
均值Mean | — | — | — | 55.77 | — | — |
表4
青杨叶片表型性状平均变异系数和Shannon-Wiener指数"
种群 Populations | 平均变异系数 Average CV(%) | 平均多样性指数 Average H | 样本数占比 Sample ratio(%) | 多样性占比 Diversity ratio(%) |
1 | 11.864±1.663 | 1.329±0.549 | 2.632 | 3.157 |
2 | 12.407±1.661 | 1.393±0.477 | 2.193 | 3.309 |
3 | 8.288±1.172 | 1.093±0.402 | 2.412 | 2.596 |
4 | 7.440±0.890 | 0.938±0.405 | 1.754 | 2.229 |
5 | 12.176±1.948 | 1.465±0.491 | 3.728 | 3.480 |
6 | 8.573±1.083 | 1.099±0.425 | 2.851 | 2.611 |
7 | 11.173±1.623 | 1.179±0.399 | 3.070 | 2.800 |
8 | 11.606±1.216 | 1.173±0.349 | 1.535 | 2.788 |
9 | 13.416±2.213 | 1.187±0.470 | 2.632 | 2.821 |
10 | 9.400±1.484 | 1.235±0.414 | 4.167 | 2.934 |
11 | 6.002±0.887 | 0.771±0.346 | 4.167 | 1.831 |
12 | 11.221±0.850 | 1.413±0.459 | 4.386 | 3.356 |
13 | 12.172±2.141 | 1.317±0.524 | 4.386 | 3.128 |
14 | 13.325±2.475 | 1.414±0.422 | 3.289 | 3.358 |
15 | 16.054±2.485 | 1.528±0.386 | 3.728 | 3.630 |
16 | 12.053±1.814 | 1.253±0.308 | 4.386 | 2.976 |
17 | 20.458±3.002 | 1.417±0.334 | 1.316 | 3.366 |
18 | 13.044±2.197 | 1.063±0.359 | 1.754 | 2.526 |
19 | 8.039±1.365 | 0.955±0.393 | 2.851 | 2.269 |
20 | 12.583±2.619 | 0.951±0.328 | 2.412 | 2.259 |
21 | 14.989±1.500 | 1.302±0.380 | 2.412 | 3.092 |
22 | 18.761±2.690 | 1.645±0.499 | 2.851 | 3.907 |
23 | 13.216±1.818 | 1.533±0.461 | 3.509 | 3.643 |
24 | 6.647±1.492 | 0.740±0.280 | 3.289 | 1.758 |
25 | 9.370±1.391 | 1.067±0.419 | 3.728 | 2.535 |
26 | 15.636±1.557 | 1.557±0.372 | 3.947 | 3.698 |
27 | 10.316±1.726 | 1.219±0.395 | 3.947 | 2.896 |
28 | 14.205±2.096 | 1.419±0.277 | 4.386 | 3.371 |
29 | 16.275±2.389 | 1.468±0.290 | 3.509 | 3.487 |
30 | 22.322±3.540 | 1.283±0.339 | 1.316 | 3.047 |
31 | 25.150±4.202 | 1.499±0.344 | 1.974 | 3.560 |
32 | 18.243±3.295 | 1.439±0.373 | 2.412 | 3.418 |
33 | 7.168±1.388 | 0.784±0.396 | 1.316 | 1.863 |
34 | 13.361±2.922 | 0.968±0.291 | 1.754 | 2.299 |
总平均Total mean | 12.852 | 1.238 | — | — |
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