青杨不同种群叶片表型性状的遗传变异

doi:10.11707/j.1001-7488.20210806

摘要/Abstract

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。

关键词: 青杨, 种群, 叶片, 遗传变异, 地理变异

Abstract:

Objective: To reveal genetic diversity and patterns of geographic variation of leaf phenotypic traits in and among different populations of Populus cathayana so as to lay a foundation for further analysis of genetic mechanism, and for collection, conservation and improvement of P. cathayana germplasm resources. Method: A total of 34 P. cathayana populations from the main distribution areas in China were sampled by collecting 1-2 years old shoots. The shoots were cutting-propagated in greenhouse and transplanted into nursery, the transplanted plants were topped in spring in the following year, the 7th, 8th and 9th new leaves below the stem tip were collected in July and 14 phenotypic traits(leaf morphological traits, leaf colour and SPAD) were measured. Variance analysis, correlation analysis and cluster analysis were conducted. Result: There were significant differences in the 14 traits among and within populations(P < 0.01), indicating that the genetic variation of leaf in P. cathayana was abundant. The average phenotypic population differentiation coefficient was 55.771% among populations, indicating the main source of variation was from among the populations. The average coefficient of variation for phenotypic traits was 20.562%, the largest coefficient of variation was relative length of petiole while the chlorophyll SPAD value was the smallest. The average coefficient of variation of phenotypic traits in each population was 12.852%, population Badi (No. 31) in Inner Mongolia Autonomous Region had the most abundant variation while population Longquan (No. 11) in Hebei province had the smallest variation and the highest stability of phenotypic traits. There was significant or extremely significant correlation between leaf phenotypic traits and geographical meteorological factors, only leaf color had no significant correlation with other leaf traits, reflecting its relatively independent inheritance. The 34 populations were divided into 4 groups, they are large leaf with long petiole, medium leaf, narrow leaf with short petiole and small leaf. The populations can be divided into three distribution regions according to the distribution of the taxa and the characteristics of Chinese topography: the northern margin of North China Plain in the east of Taihang Mountains(region 1), the Loess Plateau between Taihang Mountains and Liupan Mountains-Qinling Mountains(region 2), and the surrounding area of Qinghai-Tibet Plateau(region 3). The narrow leaf with short petiole accounted for 57.14% in the region 1, medium-sized leaf accounted for 66.67% in the region 2, and the small leaf accounted for the largest proportion 62.5% in the region 3. Conclusion: The genetic variation of leaf traits in P. cathayana was abundant, population Lanqikalun(No. 17) in Hebei province, Banjieta(No. 30) and Badi(No. 31) in Inner Mongolia Autonomous Region had rich diversity and were the priority populations for diversity protection and germplasm conservation. There was a strong correlation between leaf phenotypic traits and geographical meteorological factors, and the patterns of geographic variation of leaf of P. cathayana mainly related to altitude.

Key words: Populus cathayana, population, leaf, genetic variation, geographic variation

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.

图/表 7

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1

图1

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表4

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图3

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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

性状 Traits	方差分量Variance component			表型分化系数 Differentiation coefficients of phenotypic traits (V_st)(%)	F
性状 Traits	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	—	—

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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