Phenotypic variations in natural populations of Amygdalus pedunculata
LIUJiang-Qun, YINMing-Yu, ZUOSi-Yu, YANGShao-Bing, WUYUNTana*, Non-Timber Forest Research and Development Center, Chinese Academy of Forestry, Zhengzhou 450003, China 通讯作者:* 通信作者Author for correspondence (E-mail: tanatanan@163.ccom) 版权声明:2017植物生态学报编辑部本文是遵循CCAL协议的开放存取期刊,引用请务必标明出处。 基金资助:中央级公益性科研院所基本科研业务费专项资金(CAFYBB2017ZA004-7)
关键词:长柄扁桃;天然种群;表型变异;相关分析 Abstract Aims Our objectives were to determine the phenotypic variations, adaption and distribution patterns in seven natural Amygdalus pedunculata populations.Methods We analyzed 14 phenotypic traits from 120 individuals in seven populations of A. pedunculata by variance analysis, correlation analysis, and cluster analysis.Important findings Results showed that there were plentiful phenotypic variation within and among populations. In particular, the phenotypic variation within population was 40.91%, higher than that among populations (35.29%), which indicated that the phenotypic variation within population was the main source of the phenotypic variation in A. pedunculata. Mean differentiation coefficient was 45.90%, and mean coefficient of variation of 14 traits was 15.59%, ranged from 9.39% to 31.98%. Mean annual temperature, latitude, length of frost-free period, longitude and altitude appear to be prominent ecological factors influencing phenotypic traits. Mean annual temperature and length of frost-free period were key indicators to phenotypic of A. pedunculata in different site conditions. According to principal component analysis and unweighted pair-group method with arithmetic means (UPGMA) cluster analysis, the seven populations of A. pedunculata could be divided into two groups. In mountainous region, A. pedunculata’s leaf blade was usually rotund to oblong, fruit nearly spherical shape with shorter fruit stem, stone was usually ovoid to spherical shape. In contrast, in sandy region, leaf blade was long oval to ovate-lanceolate, fruit and stone was usually flat ovoid with longer fruit stem. Our results provide critical information for the resource collection and breeding of this ecologically important species.
FLD, fruit longitudinal diameter; FTD, fruit transverse diameter; FSD, fruit side diameter; FSI, fruit shape index; FSL, fruit stem length; LL, leaf length; LSI, leaf shape index; LW, leaf width; NLD, nutlet longitudinal diameter; NSD, nutlet side diameter; NSI, nutlet shape index; NTD, nutlet transverse diameter; PL, petiole length; PT, pulp thickness. **, p < 0.01. 新窗口打开 Table 3 表3 表3长柄扁桃7个种群的表型性状及多重比较(平均值±标准偏差) Table 3Phenotypic traits and multiple comparison of the seven Amygdalus pedunculata populations (mean ± SD)
性状 Trait
种群 Population
YYe
SYe
HJG
SSC
LMD
QD
XJG
LL (mm)
22.91 ± 4.43c
24.65 ± 5.84d
20.89 ± 3.89ab
24.15 ± 3.47cd
21.23 ± 1.99b
19.58 ± 2.28a
21.14 ± 3.172b
LW (mm)
7.70 ± 1.97a
8.19 ± 1.73a
11.41 ± 2.34b
13.09 ± 1.95c
14.63 ± 2.34d
13.75 ± 1.90c
12.15 ± 2.49b
PL (mm)
6.19 ± 1.50abc
6.96 ± 1.52d
5.95 ± 1.28ab
6.54 ± 1.31cd
5.77 ± 1.67a
6.38 ± 1.52bc
6.42 ± 1.31bc
LSI
3.11 ± 0.85c
3.06 ± 0.65c
1.86 ± 0.30b
1.86 ± 0.26b
1.48 ± 0.21a
1.44 ± 0.16a
1.80 ± 0.38b
FLD (mm)
12.81 ± 1.22b
13.35 ± 1.29c
11.79 ± 1.42a
12.62 ± 1.43b
13.59 ± 1.32c
12.14 ± 0.97a
12.74 ± 1.09b
FTD (mm)
10.11 ± 0.89a
11.32 ± 1.37c
9.98 ± 1.56a
11.00 ± 1.00bc
12.55 ± 1.31d
10.57 ± 1.03b
11.06 ± 1.29c
FSD (mm)
9.37 ± 0.85a
10.07 ± 1.00b
10.51 ± 1.47bc
11.41 ± 0.90d
12.49 ± 1.29e
10.67 ± 1.07c
11.48 ± 1.31d
FSI
1.38 ± 0.16b
1.34 ± 0.20b
1.13 ± 0.14a
1.11 ± 0.09a
1.09 ± 0.09a
1.14 ± 0.09a
1.12 ± 0.12a
FSL (mm)
6.19 ± 1.21cd
6.46 ± 1.55d
5.20 ± 1.66b
5.80 ± 1.92c
4.30 ± 1.11a
4.05 ± 1.17a
4.01 ± 1.21a
PT (mm)
1.03 ± 0.35bc
0.94 ± 0.10ab
0.86 ± 0.38a
1.16 ± 0.46c
1.15 ± 0.41c
1.46 ± 0.46d
1.40 ± 0.41d
NLD (mm)
11.88 ± 1.26b
11.74 ± 0.91b
11.03 ± 1.22a
11.46 ± 1.38a
11.13 ± 1.08a
11.00 ± 1.16a
11.64 ± 1.13b
NTD (mm)
8.48 ± 1.12a
8.98 ± 0.94b
9.38 ± 0.97cd
9.70 ± 0.72de
9.55 ± 0.56de
9.04 ± 0.95bc
9.79 ± 0.94e
NSD (mm)
7.47 ± 0.64a
7.72 ± 0.89ab
8.22 ± 1.07c
8.70 ± 0.73d
8.67 ± 0.48d
7.85 ± 0.74b
8.30 ± 0.78c
NSI
1.60 ± 0.16a
1.53 ± 0.12a
1.37 ± 0.34bc
1.32 ± 0.12ab
1.28 ± 0.11a
1.41 ± 0.13c
1.41 ± 0.17c
Table 1 and Table 2 indicated the numbers of populations and the abbreviations of phenotypic traits, respectively. Different letters after the number in the same column indicated significant difference at 0.05 level.种群缩写及表型性状缩写分别见表1和表2。表中同列数字后的不同字母表示差异显著(p < 0.05)。 新窗口打开
种群缩写及表型性状缩写分别见表1和表2。 新窗口打开 Table 5 表5 表5长柄扁桃表型性状与地理生态因子间的相关分析 Table 5Analysis of correlation between phenotypic traits and geo-ecological factors in Amygdalus pedunculata populations
性状 Trait
经度 Longitude (E)
纬度 Latitude (N)
海拔 Altitude (m)
年平均气温 AAT (℃)
年日照时间 AAS (h)
年降水量 AP (mm)
无霜期 Frost-free season (d)
LL
-0.344
-0.593
-0.439
0.661
-0.095
0.189
0.606
LW
0.455
0.910**
0.356
-0.958**
0.625
-0.426
-0.797*
PL
0.022
-0.365
0.134
0.513
-0.123
0.377
0.416
LSI
-0.521
-0.959**
-0.478
0.982**
-0.573
0.436
0.862*
FLD
0.001
-0.345
-0.851*
0.269
-0.399
0.537
0.336
FTD
0.292
0.229
-0.566
-0.281
-0.026
0.223
-0.123
FSD
0.594
0.779*
-0.075
-0.810*
0.336
-0.118
-0.707
FSL
-0.676
-0.787*
-0.347
0.842*
-0.119
-0.006
0.813*
PT
0.555
0.493
0.471
-0.566
0.149
0.243
-0.594
FSI
-0.600
-0.988**
-0.419
0.971**
-0.586
0.430
0.899**
NLD
-0.086
-0.729
-0.436
0.732
-0.610
0.721
0.494
NTD
0.778*
0.845*
0.187
-0.755*
0.337
-0.116
-0.801*
NSD
0.486
0.788*
0.043
-0.772*
0.531
-0.354
-0.706
NSI
-0.417
-0.893**
-0.218
0.889**
-0.656
0.529
0.739
Total
5.827
9.701
5.019
10.003
5.165
4.705
8.894
Abbreviations see Table 1 and Table 2. * and ** indicate significant correlation at p < 0.05 and p < 0.01, respectively.缩写同表1、表2。*与**分别表示在p < 0.05和p < 0.01的显著水平。 新窗口打开
Abbreviations of phenotypic traits was indicated in Table 2.表型性状缩写同表2。 新窗口打开 显示原图|下载原图ZIP|生成PPT 图1基于表型性状(PC-1, PC-2)的长柄扁桃种群关系。种群缩写见表1。 -->Fig. 1The relationship among different Amygdalus pedunculata populations based on PC-1 and PC-2. Table 1 indicated the numbers of populations. -->
2.6 种群聚类分析及不同立地条件下的长柄扁桃表型变异
对7个种群进行UPGMA聚类分析(图2), 基于表型性状将种群划分为两大类, 第一大类包括HJG、XJG、QD、LMD和SSC, 主要立地为山地; 第二大类包括YYe和SYe, 主要立地为沙地。通过比较山地、沙地这两种不同立地条件下生长的长柄扁桃表型性状(表7), 可知立地条件对长柄扁桃不同表型性状具有不同程度的影响, 其中, 对叶长、叶宽、叶形指数、果侧径、果柄长、肉厚、果形指数、核横径、核侧径及核形指数等性状的影响较大。相对于山地上生长的长柄扁桃, 沙地的长柄扁桃叶更长更窄、叶形指数更大, 山地的长柄扁桃叶形为近圆形至长圆形, 而沙地的多为长椭圆形或卵状披针形(图3); 沙地的长柄扁桃的果纵径、果柄、果形指数都大于山地的长柄扁桃, 果横径小于山地的长柄扁桃, 沙地的果实多为卵球形, 而山地的果实多为近球形, 沙地的长柄扁桃果柄普遍长于山地(图4); 沙地的长柄扁桃核横径和核侧径均小于山地的长柄扁桃, 核形指数大于山地的长柄扁桃, 沙地的长柄扁桃核近宽卵形, 山地的长柄扁桃核近宽卵形至近球形。不同立地的长柄扁桃表型特征差异对比见表8。 显示原图|下载原图ZIP|生成PPT 图2基于表型性状的长柄扁桃种群聚类分析。种群缩写同表1。 -->Fig. 2Cluster analysis based on the phenotypic traits of Amygdalus pedunculata populations. Table 1 indicated the numbers of populations. -->
显示原图|下载原图ZIP|生成PPT 图3不同立地下长柄扁桃叶片形态变异。A, 山地。B, 沙地。网格大小为1 cm × 1 cm。 -->Fig. 3The phenotype difference of leaves in Amygdalus pedunculata between different stands. A, Mountainous region. B, Sand. The size of the grid was 1 cm × 1 cm. -->
显示原图|下载原图ZIP|生成PPT 图4不同立地下长柄扁桃果实形态变异。A, 山地果枝。B, 山地果实。C, 山地果肉。D, 沙地果枝。E, 沙地果实。F, 沙地果肉。网格大小为1 cm × 1 cm。 -->Fig. 4The phenotype difference of fruits in Amygdalus pedunculata between different stands. A, Fruit spur from mountainous region. B, Fruit from mountainous region. C, Pulp from mountainous region. D, Fruit spur from sand. E, Fruit from sand. F, Pulp from sand. The size of the grid was 1 cm × 1 cm. -->
Table 7 表7 表7不同立地对长柄扁桃表型的影响 Table 7The influence of different stands on Amygdalus pedunculata populations
性状 Trait
山地 Mountainous region
沙地 Sand
t检验 t-test
分布范围 Rangeability
均值 Mean value
标准偏差 Standard deviation
分布范围 Rangeability
均值 Mean value
标准偏差 Standard deviation
LL (mm)
9.21-33.19
20.75
4.04
12.16-38.91
23.45
4.96
**
LW (mm)
5.21-19.52
12.01
2.61
4.23-13.95
7.85
1.90
**
PL (mm)
2.68-12.22
6.28
1.43
2.87-10.44
6.44
1.57
-
LSI
0.84-3.23
1.77
0.35
1.96-5.93
3.08
0.72
**
FLD (mm)
7.14-19.22
12.65
1.45
9.75-16.21
12.98
1.26
*
FTD (mm)
5.58-14.77
10.68
1.39
8.30-14.65
10.49
1.19
-
FSD (mm)
5.92-15.73
11.20
1.33
7.24-12.06
9.59
0.95
**
FSL (mm)
1.16-10.15
4.47
1.54
3.08-10.07
6.27
1.32
**
PT (mm)
0.15-2.99
1.23
0.47
0.38-2.21
1.00
0.30
**
FSI
0.75-1.66
1.14
0.14
0.94-1.82
1.36
0.17
**
NLD (mm)
7.28-18.15
11.61
1.44
9.89-15.27
11.84
1.16
-
NTD (mm)
7.04-13.45
9.67
0.97
5.82-10.97
8.64
1.09
**
NSD (mm)
2.48-11.93
8.27
0.85
6.09-9.64
7.54
0.73
**
NSI
0.81-4.06
1.42
0.25
1.23-2.01
1.58
0.15
**
Abbreviations of phenotypic traits was indicated in Table 2. * and ** indicate significant correlation at p < 0.05 and p < 0.01, respectively.表型性状缩写同表2。*与**分别表示在p < 0.05和p < 0.01的显著水平。 新窗口打开 Table 8 表8 表8不同立地长柄扁桃形态特征对比 Table 8Comparisons of morphological characteristics of Amygdalus pedunculata populations between different sites
山地 Mountainous region
沙地 Sand
叶片形态 Leaf morphology
叶片近圆形至长圆形, 长9.21-33.19 mm, 宽5.21-19.52 mm; 叶柄长2.68-12.22 mm。 Blade rotund to oblong, 9.21-33.19 mm long, 5.21-19.52 mm wide; petioles 2.68-12.22 mm long.
叶长椭圆形或卵状披针形, 长12.16-38.91 mm, 宽4.23-13.95 mm; 叶柄长2.87-10.44 mm。 Blade long oval to ovate-lanceolate, 12.16-38.91 mm long, 4.23- 13.95 mm wide; petioles 2.87-10.44 mm long.
果实形态 Fruit morphology
果实近球形, 直径5.92-15.73 mm; 果柄较短, 长1.16-10.15 mm; 果肉较厚, 0.15-2.99 mm。 Fruit subglobose, 5.92-15.73 mm in diameter; fruit stem short, 1.16-10.15 mm; pulp thick, 0.15-2.99 mm.
果实卵球形, 直径7.24-12.06 mm; 果柄较长, 长3.08-10.07 mm; 果肉较薄, 0.38-2.21 mm。 Fruit ovoid, 7.24-12.06 mm in diameter; fruit stem long, 3.08-10.07 mm; pulp thin, 0.38-2.21 mm.
核形态 Stone form
核宽卵形至近球形, 直径2.48-11.93 mm。 Stone wide oval to subglobose, 2.48-11.93 mm in diameter.
核宽卵形, 直径6.09-9.64 mm。 Stone wide oval to subglobose, 6.09-9.64 mm in diameter.
新窗口打开 对不同立地条件各地理生态因子进行差异显著性检验(表9), 发现年平均气温和无霜期差异均极显著(p < 0.01), 纬度差异显著(p < 0.05), 表明年平均气温、无霜期是不同立地长柄扁桃表型差异的主要影响因子。 Table 9 表9 表9不同立地各地理生态因子的比较分析 Table 9Comparisons on geographical and ecological factors between different sites
地理生态因子 Geo-ecological factor
山地 Mountainous region
沙地 Sand
t检验 t-test
均值 Mean value
标准偏差 Standard deviation
均值 Mean value
标准偏差 Standard deviation
经度 Longitude (E)
110.48
0.76
109.535
0.47
-
纬度 Latitude (N)
40.81
0.10
38.65
0.28
*
海拔 Altitude (m)
1 394.00
107.60
1 281.00
18.38
-
年平均气温 AAT (℃)
3.78
0.52
8.45
0.49
**
年日照时间 AAS (h)
2 974.80
78.85
2 877.50
2.12
-
年降水量 AP (mm)
352.58
82.62
425.25
16.05
-
无霜期 Frost-free season (d)
113.80
10.64
161.50
10.60
**
AAT, AAS, AP see Table 1. * and ** indicate significant correlation at p < 0.05 and p < 0.01, respectively.AAT、AAS和AP同表1。*与**分别表示在p < 0.05和p < 0.01的显著水平。 新窗口打开
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