东北杏种质资源多样性及其地理变化

doi:10.17521/cjpe.2019.0060

摘要/Abstract

摘要：

东北杏(Armeniaca mandshurica)是集观赏、经济、用材于一体的重要树种, 长期以来主要处于野生和半野生状态, 鲜有相关研究报道。该研究对东北杏主要分布区内种质资源状况开展了调查, 选择来自辽宁、吉林与黑龙江的47份典型种质进行了22个定量描述性状和7个定性描述性状的观测, 旨在为该树种的种质资源收集、评价与保护提供重要参考。采用变异系数等指标分析定量描述性状多样性, 使用频率分布等指标分析定性描述性状多样性。通过趋势面方法分析定量描述性状的地理变化规律。利用99对SSR引物对47份东北杏种质进行PCR扩增, 应用遗传相似系数分析东北杏种质的遗传多样性。基于表型性状和SSR分子标记采用聚类分析方法分别对种质进行分类。东北杏不同种质间呈现出较高的表型多样性, 其中19个定量描述性状指标的变异系数在9.40%-55.98%之间, 变异系数最大的为小枝长度, 变异系数最小的为种仁宽; 7个定性描述性状的Shannon-Wiener指数在0.58-1.22之间。由于调查区域的地理位置与主要气候因子间存在着显著的相关性, 东北杏种质的定量描述性状与其地理位置有着密切的关系。其中, 小枝长度呈由东向西逐渐增大的梯度变化, 小枝粗度、种子质量呈由北向南逐渐增大的梯度变化, 果柄长呈由东北向西南逐渐增大的梯度变化; 小枝长度、果柄长与海拔呈正相关关系, 小枝粗度与海拔呈负相关关系, 种子质量与海拔相关性很小。基于26个表型性状进行系统聚类, 将47份东北杏种质分为4类, 分类结果主要体现了东北杏种质特征的差别, 同时也在一定程度上体现了种质产地的效应; 基于遗传相似系数进行聚类分析, 将47份东北杏种质也分为4类, 分类结果体现了种质产地效应; 卡方检验表明, 两种聚类结果相关性不显著, 外在环境是影响东北杏表型变异的重要因素。

关键词: 东北杏, 多样性, 表型性状, 地理变化, SSR标记

Abstract:

Aims Armeniaca mandshurica is an important species which serves the need of ornamental, wood and other economical uses. This species has been in wild or semi-wild state for a long time and few studies about this species have been conducted. This paper aimed to provide an important reference for the collection, evaluation and protection of germplasm resources of A. mandshurica.
Methods The status of the germplasm resources in the main distribution area of A. mandshurica was investigated. In total, 47 typical sample trees from Liaoning, Jilin and Heilongjiang Provinces were selected. For each tree, 22 quantitative traits and 7 qualitative traits were measured. The diversity of quantitative traits was represented by indicators such as coefficient of variation, and the diversity of qualitative traits was represented by indicators such as frequency distribution. Trend surface analysis was used to explore the geographical variation in the quantitative traits. A total of 115 pairs of SSR primers were used for PCR amplification for 47 A. mandshurica germplasms, and the genetic diversity of A. mandshurica germplasms was analyzed using genetic similarity coefficient. Germplasms were classified by cluster analysis based on phenotypic traits and SSR markers separately.
Important findings High phenotypic diversity was found among different germplasms in A. mandshurica. The coefficients of variation (CV) of the 19 quantitative traits ranged from 9.40% to 55.98%. Among the 19 traits, twig length had the highest CV and kernel width had the lowest CV. The Shannon-Wiener index of 7 qualitative traits ranged from 0.58 to 1.22. Due to the significant correlation between geographical locations and main climatic factors within the study area, the quantitative traits of A. mandshurica germplasms were closely related to their geographical locations. Among them, twig length increased from east to west, twig thickness and seed mass increased from north to south, and fruit handle length increased from northeast to southwest. Twig length and fruit handle length were positively correlated with altitude, twig thickness was negatively correlated with altitude, and seed mass was not correlated with altitude. Clustering analysis based on 26 phenotypic traits showed that the 47 A. mandshurica germplasms could be divided into 4 categories, which mainly reflected the difference of the germplasm characteristics in A. mandshurica and to some extent also reflected the difference of germplasm productions. Clustering analysis based on genetic similarity coefficients showed that the 47 A. mandshurica germplasms were also divided into 4 categories, which reflected the difference of germplasm productions. Chi-square test showed that the correlation between the two clustering results was not significant, and that the external environment was the key factor affecting phenotypic variations in A. mandshurica.

Key words: Armeniaca mandshurica, diversity, phenotypic traits, geographical variations, SSR markers

徐豪, 刘明国, 董胜君, 吴月亮, 张皓凯. 东北杏种质资源多样性及其地理变化. 植物生态学报, 2019, 43(7): 585-600. DOI: 10.17521/cjpe.2019.0060

XU Hao, LIU Ming-Guo, DONG Sheng-Jun, WU Yue-Liang, ZHANG Hao-Kai. Diversity and geographical variations of germplasm resources of Armeniaca mandshurica. Chinese Journal of Plant Ecology, 2019, 43(7): 585-600. DOI: 10.17521/cjpe.2019.0060

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2019.0060

https://www.plant-ecology.com/CN/Y2019/V43/I7/585

图/表 14

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种质编号 Germplasm No.	地点 Site	海拔 Altitude (m)	经纬度 Longitude (E) and Latitude (N)	种质编号 Germplasm No.	地点 Site	海拔 Altitude (m)	经纬度 Longitude (E) and Latitude (N)
701	凤城 Fengcheng	185.6	123.95°, 40.42°	747	桦甸 Huadian	268.5	127.04°, 42.98°
702	凤城 Fengcheng	187.6	123.95°, 40.42°	748	敦化 Dunhua	495.0	128.22°, 43.38°
703	凤城 Fengcheng	185.4	123.95°, 40.42°	749	敦化 Dunhua	509.6	128.22°, 43.38°
704	本溪 Benxi	406.0	123.96°, 40.88°	750	敦化 Dunhua	517.3	128.22°, 43.38°
705	本溪 Benxi	405.8	123.96°, 40.88°	751	敦化 Dunhua	598.8	128.54°, 43.41°
706	本溪 Benxi	406.3	123.96°, 40.88°	752	敦化 Dunhua	599.1	128.54°, 43.41°
707	桓仁 Huanren	277.6	125.37°, 41.25°	753	敦化 Dunhua	595.3	128.54°, 43.41°
708	桓仁 Huanren	272.6	125.37°, 41.25°	755	吉林 Jilin	301.4	126.80°, 43.97°
709	桓仁 Huanren	283.3	125.37°, 41.25°	756	吉林 Jilin	302.2	126.80°, 43.97°
710	新宾 Xinbin	387.1	125.10°, 41.77°	757	吉林 Jilin	303.7	126.80°, 43.97°
711	新宾 Xinbin	416.5	125.10°, 41.77°	758	吉林 Jilin	200.0	126.55°, 43.85°
712	新宾 Xinbin	419.1	125.10°, 41.77°	772	宁安 Ningan	263.8	129.52°, 44.42°
713	清原 Qingyuan	315.7	124.81°, 42.34°	773	宁安 Ningan	262.2	129.52°, 44.42°
714	清原 Qingyuan	318.7	124.81°, 42.34°	774	宁安 Ningan	268.6	129.52°, 44.42°
715	清原 Qingyuan	315.7	124.81°, 42.34°	775	宁安 Ningan	279.6	129.52°, 44.42°
724	沈阳 Shenyang	60.0	123.57°, 41.82°	776	尚志 Shangzhi	327.6	127.55°, 45.27°
729	沈阳 Shenyang	58.0	123.55°, 41.82°	777	鸡西 Jixi	303.6	130.52°, 45.21°
730	沈阳 Shenyang	65.0	123.55°, 41.82°	778	鸡西 Jixi	282.6	130.54°, 45.18°
741	抚松 Fusong	461.9	127.20°, 42.34°	779	鸡西 Jixi	372.4	130.69°, 45.02°
742	抚松 Fusong	460.9	127.20°, 42.34°	783	阿城 Acheng	243.0	127.05°, 45.44°
743	抚松 Fusong	449.7	127.20°, 42.34°	784	阿城 Acheng	262.0	127.05°, 45.43°
744	磐石 Panshi	399.2	126.06°, 42.86°	785	阿城 Acheng	262.1	127.05°, 45.43°
745	磐石 Panshi	402.5	126.06°, 42.86°	786	阿城 Acheng	216.4	127.06°, 45.44°
746	磐石 Panshi	401.3	126.06°, 42.86°

种质编号 Germplasm No.	地点 Site	海拔 Altitude (m)	经纬度 Longitude (E) and Latitude (N)	种质编号 Germplasm No.	地点 Site	海拔 Altitude (m)	经纬度 Longitude (E) and Latitude (N)
701	凤城 Fengcheng	185.6	123.95°, 40.42°	747	桦甸 Huadian	268.5	127.04°, 42.98°
702	凤城 Fengcheng	187.6	123.95°, 40.42°	748	敦化 Dunhua	495.0	128.22°, 43.38°
703	凤城 Fengcheng	185.4	123.95°, 40.42°	749	敦化 Dunhua	509.6	128.22°, 43.38°
704	本溪 Benxi	406.0	123.96°, 40.88°	750	敦化 Dunhua	517.3	128.22°, 43.38°
705	本溪 Benxi	405.8	123.96°, 40.88°	751	敦化 Dunhua	598.8	128.54°, 43.41°
706	本溪 Benxi	406.3	123.96°, 40.88°	752	敦化 Dunhua	599.1	128.54°, 43.41°
707	桓仁 Huanren	277.6	125.37°, 41.25°	753	敦化 Dunhua	595.3	128.54°, 43.41°
708	桓仁 Huanren	272.6	125.37°, 41.25°	755	吉林 Jilin	301.4	126.80°, 43.97°
709	桓仁 Huanren	283.3	125.37°, 41.25°	756	吉林 Jilin	302.2	126.80°, 43.97°
710	新宾 Xinbin	387.1	125.10°, 41.77°	757	吉林 Jilin	303.7	126.80°, 43.97°
711	新宾 Xinbin	416.5	125.10°, 41.77°	758	吉林 Jilin	200.0	126.55°, 43.85°
712	新宾 Xinbin	419.1	125.10°, 41.77°	772	宁安 Ningan	263.8	129.52°, 44.42°
713	清原 Qingyuan	315.7	124.81°, 42.34°	773	宁安 Ningan	262.2	129.52°, 44.42°
714	清原 Qingyuan	318.7	124.81°, 42.34°	774	宁安 Ningan	268.6	129.52°, 44.42°
715	清原 Qingyuan	315.7	124.81°, 42.34°	775	宁安 Ningan	279.6	129.52°, 44.42°
724	沈阳 Shenyang	60.0	123.57°, 41.82°	776	尚志 Shangzhi	327.6	127.55°, 45.27°
729	沈阳 Shenyang	58.0	123.55°, 41.82°	777	鸡西 Jixi	303.6	130.52°, 45.21°
730	沈阳 Shenyang	65.0	123.55°, 41.82°	778	鸡西 Jixi	282.6	130.54°, 45.18°
741	抚松 Fusong	461.9	127.20°, 42.34°	779	鸡西 Jixi	372.4	130.69°, 45.02°
742	抚松 Fusong	460.9	127.20°, 42.34°	783	阿城 Acheng	243.0	127.05°, 45.44°
743	抚松 Fusong	449.7	127.20°, 42.34°	784	阿城 Acheng	262.0	127.05°, 45.43°
744	磐石 Panshi	399.2	126.06°, 42.86°	785	阿城 Acheng	262.1	127.05°, 45.43°
745	磐石 Panshi	402.5	126.06°, 42.86°	786	阿城 Acheng	216.4	127.06°, 45.44°
746	磐石 Panshi	401.3	126.06°, 42.86°

序号 No.	性状 Traits	表型性状描述 Description of phenotypic traits
1	小枝色泽 Twig color	(1)绿色 Green (2)黄褐色 Tawny (3)灰褐色 Grayish-brown
2	叶背被毛 Leaf back fuzzed	(1)有 Hairy (2)无 Glabrous
3	叶表被毛 Leaf surface fuzzed	(1)有 Hairy (2)无 Glabrous
4	叶基形状 Leaf base shape	(1)宽楔形 Wide wedge (2)圆形 Round (3)窄楔形 Narrow wedge
5	叶缘形状 Leaf edge shape	(1)锐重锯齿 Sharp double sawtooth (2)钝重锯齿 Blunt double sawtooth
6	腺体数量 Number of glands	(1) 1个 One (2) 2个 Two (3)无 Zero
7	果实形状 Fruit shape	(1)球形 Sphericity (2)扁球形 Flat sphericity (3)扁长球形 Prolate sphericity (4)扁宽球形 Flat and broad sphericity (5)长球形 Long sphericity
8	树高 Tree height	树木从根颈到树梢之间的距离 The distance between the treetop and the root neck of tree
9	胸径 Diameter at breast height	树干1.3 m高度处的直径 Diameter of the trunk at 1.3 m height
10	冠幅 Crown diameter	树冠南北和东西方向宽度的平均值 The average value of the north-south and east-west width of tree crown
11	主枝基角 Main branch base angle	主枝与中心干之间的分枝角度 Angle between the main branch and the center stem of tree
12	小枝长度 Twig length	一年生枝平均长度 Average length of annual branches
13	小枝粗度 Twig width	一年生枝平均粗度 Average width of annual branches
14	叶长 Leaf length	从叶基切线至叶尖顶部的长度 Length from the tangent of leaf base to the top of leaf tip
15	叶宽 Leaf width	叶片最宽处的长度 The length of the blade at its widest point
16	叶柄长 Petiole length	叶柄的长度 Length of petiole
17	单果质量 Single fruit mass	单个成熟果实的质量 The mass of single ripe fruit
18	果长 Fruit length	成熟果实从顶部到底部的最大距离 Maximum distance from top to bottom of ripe fruit
19	果宽 Fruit width	成熟果实两条缝合线之间的最大距离 Maximum distance between two sutures of ripe fruit
20	果厚 Fruit thickness	成熟果实腹面观时的最大距离 Maximum distance from the ventral view of ripe fruit
21	果柄长 Fruit handle length	果柄的长度 Length of fruit handle
22	种子质量 Seed mass	单个成熟种子的质量 The mass of single ripe seed
23	种子长 Seed length	成熟种子从顶部到底部的最大距离 Maximum distance from top to bottom of ripe seed
24	种子宽 Seed width	成熟种子脊背到缝翅的最大距离 Maximum distance from ridge to sewed wing of ripe seed
25	种子厚 Seed thickness	成熟种子腹面观时的最大距离 Maximum distance from the ventral view of ripe seed
26	种仁质量 Kernel mass	单个成熟种仁的质量 The mass of single mature kernel
27	种仁长 Kernel length	成熟种仁从顶部到底部的最大距离 Maximum distance from top to bottom of ripe kernel
28	种仁宽 Kernel width	成熟种仁两条缝合线之间的最大距离 Maximum distance between two sutures of ripe kernel
29	种仁厚 Kernel thickness	成熟种仁腹面观时的最大距离 Maximum distance from the ventral view of ripe kernel

序号 No.	性状 Traits	表型性状描述 Description of phenotypic traits
1	小枝色泽 Twig color	(1)绿色 Green (2)黄褐色 Tawny (3)灰褐色 Grayish-brown
2	叶背被毛 Leaf back fuzzed	(1)有 Hairy (2)无 Glabrous
3	叶表被毛 Leaf surface fuzzed	(1)有 Hairy (2)无 Glabrous
4	叶基形状 Leaf base shape	(1)宽楔形 Wide wedge (2)圆形 Round (3)窄楔形 Narrow wedge
5	叶缘形状 Leaf edge shape	(1)锐重锯齿 Sharp double sawtooth (2)钝重锯齿 Blunt double sawtooth
6	腺体数量 Number of glands	(1) 1个 One (2) 2个 Two (3)无 Zero
7	果实形状 Fruit shape	(1)球形 Sphericity (2)扁球形 Flat sphericity (3)扁长球形 Prolate sphericity (4)扁宽球形 Flat and broad sphericity (5)长球形 Long sphericity
8	树高 Tree height	树木从根颈到树梢之间的距离 The distance between the treetop and the root neck of tree
9	胸径 Diameter at breast height	树干1.3 m高度处的直径 Diameter of the trunk at 1.3 m height
10	冠幅 Crown diameter	树冠南北和东西方向宽度的平均值 The average value of the north-south and east-west width of tree crown
11	主枝基角 Main branch base angle	主枝与中心干之间的分枝角度 Angle between the main branch and the center stem of tree
12	小枝长度 Twig length	一年生枝平均长度 Average length of annual branches
13	小枝粗度 Twig width	一年生枝平均粗度 Average width of annual branches
14	叶长 Leaf length	从叶基切线至叶尖顶部的长度 Length from the tangent of leaf base to the top of leaf tip
15	叶宽 Leaf width	叶片最宽处的长度 The length of the blade at its widest point
16	叶柄长 Petiole length	叶柄的长度 Length of petiole
17	单果质量 Single fruit mass	单个成熟果实的质量 The mass of single ripe fruit
18	果长 Fruit length	成熟果实从顶部到底部的最大距离 Maximum distance from top to bottom of ripe fruit
19	果宽 Fruit width	成熟果实两条缝合线之间的最大距离 Maximum distance between two sutures of ripe fruit
20	果厚 Fruit thickness	成熟果实腹面观时的最大距离 Maximum distance from the ventral view of ripe fruit
21	果柄长 Fruit handle length	果柄的长度 Length of fruit handle
22	种子质量 Seed mass	单个成熟种子的质量 The mass of single ripe seed
23	种子长 Seed length	成熟种子从顶部到底部的最大距离 Maximum distance from top to bottom of ripe seed
24	种子宽 Seed width	成熟种子脊背到缝翅的最大距离 Maximum distance from ridge to sewed wing of ripe seed
25	种子厚 Seed thickness	成熟种子腹面观时的最大距离 Maximum distance from the ventral view of ripe seed
26	种仁质量 Kernel mass	单个成熟种仁的质量 The mass of single mature kernel
27	种仁长 Kernel length	成熟种仁从顶部到底部的最大距离 Maximum distance from top to bottom of ripe kernel
28	种仁宽 Kernel width	成熟种仁两条缝合线之间的最大距离 Maximum distance between two sutures of ripe kernel
29	种仁厚 Kernel thickness	成熟种仁腹面观时的最大距离 Maximum distance from the ventral view of ripe kernel

序号 No.	性状 Traits	频率分布 Frequency distribution (%)					Shannon-Wiener 多样性指数 Shannon-Wiener diversity index
序号 No.	性状 Traits	1	2	3	4	5	Shannon-Wiener 多样性指数 Shannon-Wiener diversity index
1	小枝色泽 Twig color	80.85	14.89	4.26			0.59
2	叶背被毛 Leaf back fuzzed	74.47	25.53				0.57
3	叶表被毛 Leaf surface fuzzed	68.09	31.91				0.63
4	叶基形状 Leaf base shape	74.47	23.40	2.13			0.64
5	叶缘形状 Leaf edge shape	51.06	48.94				0.69
6	腺体数量 Number of glands	34.04	40.43	25.53			1.08
7	果实形状 Fruit shape	23.41	53.19	14.89	6.38	2.13	1.22