Table 1. Characteristics of physical and chemical analysis of the studied soil
Organic matter %
|
E.C. (ds/m)
|
pH
|
Total lime %
|
Clay %
|
Silt %
|
Sand %
|
N %
|
Total P (mg/kg)
|
Total K (mg/kg)
|
Total Mg (meq/l)
|
Total Ca (meq/l)
|
Soil texture
|
1.18
|
0.72
|
8.28
|
7.2
|
33
|
27
|
40
|
0.09
|
9.6
|
286
|
1.1
|
2.1
|
Loamy-clay
|
Table 2. Variance analysis table
Source of Variation
|
df
|
Expected value
|
Block
|
r-1
|
|
Population
|
p-1
|
σ2e +r σ2t
|
Error
|
(r-1)(p-1)
|
σ2e
|
Table 3. Variability of phenotypic and genotypic parameters among Glycyrrhiza glabra L. populations.
Traits
|
VP
|
VG
|
VA
|
PCV
|
GCV
|
H2b
|
H2n
|
GA
|
GAM
|
Plant height
|
413.70
|
344.25
|
86.06
|
33.75
|
30.78
|
83.21
|
20.80
|
1.88
|
3.12
|
Plant width
|
401.34
|
235.89
|
58.97
|
29.82
|
22.86
|
58.78
|
14.69
|
1.58
|
2.35
|
Main stem diameter
|
4.05
|
1.57
|
0.39
|
28.36
|
17.67
|
38.83
|
9.71
|
1.28
|
18.08
|
Leaf length
|
3.46
|
2.05
|
0.51
|
22.86
|
17.59
|
59.17
|
14.79
|
1.58
|
19.47
|
Leaf width
|
16.32
|
7.83
|
1.96
|
28.09
|
19.46
|
47.99
|
12.00
|
1.43
|
9.92
|
Number of leaflets
|
1.88
|
1.33
|
0.33
|
11.69
|
9.85
|
71.06
|
17.77
|
1.74
|
14.82
|
Leaflet length
|
42.00
|
28.09
|
7.02
|
25.97
|
21.24
|
66.88
|
16.72
|
1.68
|
6.75
|
Leaflet width
|
3.10
|
1.50
|
0.37
|
26.08
|
18.12
|
48.24
|
12.06
|
1.43
|
21.20
|
No. of axillary branches
|
1.94
|
1.04
|
0.26
|
29.41
|
21.47
|
53.32
|
13.33
|
1.50
|
31.73
|
Shoot fresh weight
|
17280.93
|
8019.77
|
2004.94
|
52.26
|
35.60
|
46.41
|
11.60
|
1.40
|
0.56
|
Shoot dry weight
|
5270.98
|
2313.50
|
578.38
|
52.53
|
34.80
|
43.89
|
10.97
|
1.36
|
0.99
|
Shoot yield
|
51474.51
|
22592.86
|
5648.22
|
52.52
|
34.80
|
43.89
|
10.97
|
1.36
|
0.32
|
Root fresh weight
|
15801.12
|
6149.93
|
1537.48
|
38.95
|
24.30
|
38.92
|
9.73
|
1.29
|
0.40
|
Root dry weight
|
7278.80
|
2977.39
|
744.35
|
42.42
|
27.13
|
40.90
|
10.23
|
1.32
|
0.66
|
Root yield
|
71081.74
|
29075.90
|
7268.98
|
42.42
|
27.13
|
97.76
|
24.44
|
1.37
|
0.32
|
Glycyrrhizic acid
|
1561.94
|
1539.53
|
384.88
|
40.90
|
40.60
|
98.56
|
24.64
|
2.05
|
2.12
|
Glabridin
|
19.66
|
19.51
|
4.88
|
156.68
|
156.07
|
99.22
|
24.80
|
2.05
|
72.51
|
Liquritin
|
1.72
|
1.67
|
0.42
|
76.82
|
75.67
|
97.03
|
24.26
|
2.03
|
118.88
|
Liquritigenin
|
9.47
|
9.28
|
2.32
|
46.08
|
45.61
|
97.96
|
24.49
|
2.04
|
30.52
|
TPC
|
2.43
|
1.71
|
0.43
|
25.37
|
21.25
|
70.19
|
17.55
|
1.73
|
28.09
|
TFC
|
34.28
|
16.72
|
4.18
|
35.96
|
25.11
|
48.78
|
12.20
|
1.44
|
8.84
|
IC50
|
51.02
|
24.98
|
6.25
|
8.93
|
6.25
|
48.97
|
12.24
|
1.44
|
1.80
|
Rutin
|
0.58
|
0.58
|
0.15
|
20.37
|
20.35
|
99.79
|
24.95
|
2.06
|
54.98
|
Rosmarinic acid
|
0.03
|
0.03
|
0.01
|
35.01
|
34.84
|
99.00
|
24.75
|
2.05
|
414.91
|
VG: Genotypic variance, VP: Phenotypic variance, VA: Additive variance, H2b: Broad sense heritability, H2n: Narrow sense heritability, PCV: Phenotypic coefficient of variation, GCV: Genotypic coefficient of variation, GA: Genetic advance, GAM: Genetic advance as percentage of mean.
Table 4. Phenotypic (P) and genotypic (G) correlation coefficients of measured traits in G. glabra
Correlations
|
|
|
PH
|
PW
|
MSD
|
LL
|
LW
|
NL
|
LEL
|
LEW
|
NAB
|
SFW
|
SDW
|
SY
|
|
PH
|
P
|
1
|
|
|
|
|
|
|
|
|
|
|
|
|
G
|
1
|
|
|
|
|
|
|
|
|
|
|
|
|
PW
|
P
|
0.644**
|
1
|
|
|
|
|
|
|
|
|
|
|
|
G
|
0.778
|
1
|
|
|
|
|
|
|
|
|
|
|
|
MSD
|
P
|
0.652**
|
0.546**
|
1
|
|
|
|
|
|
|
|
|
|
|
G
|
0.895
|
0.875
|
1
|
|
|
|
|
|
|
|
|
|
|
LL
|
P
|
0.553**
|
0.361**
|
0.565**
|
1
|
|
|
|
|
|
|
|
|
|
G
|
0.686
|
0.432
|
0.984
|
1
|
|
|
|
|
|
|
|
|
|
LW
|
P
|
0.194*
|
0.075
|
0.322**
|
0.321**
|
1
|
|
|
|
|
|
|
|
|
G
|
0.312
|
0.177
|
0.821
|
0.780
|
1
|
|
|
|
|
|
|
|
|
NL
|
P
|
0.345**
|
0.292**
|
0.271**
|
0.433**
|
-0.198*
|
1
|
|
|
|
|
|
|
|
G
|
0.413
|
0.439
|
0.940
|
0.893
|
0.825
|
1
|
|
|
|
|
|
|
|
LEL
|
P
|
0.404**
|
0.278**
|
0.438**
|
0.407**
|
0.657**
|
-0.130
|
1
|
|
|
|
|
|
|
G
|
0.627
|
0.595
|
0.850
|
0.625
|
0.915
|
-0.069
|
1
|
|
|
|
|
|
|
LEW
|
P
|
0.194*
|
0.075
|
0.322**
|
0.321**
|
0.884**
|
-0.198*
|
0.657**
|
1
|
|
|
|
|
|
G
|
0.269
|
0.125
|
0.998
|
0.782
|
0.995
|
0.089
|
0.915
|
1
|
|
|
|
|
|
NAB
|
P
|
.598**
|
0.631**
|
0.435**
|
0.446**
|
0.318**
|
0.093
|
0.461**
|
0.318**
|
1
|
|
|
|
|
G
|
0.778
|
0.711
|
1.473
|
0.863
|
0.743
|
0.630
|
0.873
|
0.759
|
1
|
|
|
|
|
SFW
|
P
|
0.506**
|
0.621**
|
0.459**
|
0.244*
|
0.245**
|
-0.023
|
0.443**
|
0.245**
|
0.620**
|
1
|
|
|
|
G
|
0.997
|
0.894
|
0.798
|
0.409
|
0.359
|
-0.012
|
0.784
|
0.350
|
0.727
|
1
|
|
|
|
SDW
|
P
|
0.485**
|
0.564**
|
0.472**
|
0.264**
|
0.234*
|
-0.018
|
0.400**
|
0.234*
|
0.641**
|
0.942**
|
1
|
|
|
G
|
0.984
|
0.943
|
-0.650
|
0.491
|
0.358
|
0.045
|
0.792
|
0.341
|
0.750
|
0.982
|
1
|
|
|
SY
|
P
|
0.495**
|
0.562**
|
0.491**
|
0.251**
|
0.248*
|
-0.032
|
0.425**
|
0.244*
|
0.637**
|
0.969**
|
0.989**
|
1
|
|
G
|
0.953
|
0.992
|
0.826
|
0.481
|
0.338
|
-0.037
|
0.778
|
0.322
|
0.774
|
1.000
|
1.000
|
1
|
|
Table 4 Continued. Phenotypic (P) and genotypic (G) correlation coefficients of measured traits in G. glabra
Genotypic (G) and Phenotypic Correlations
|
|
|
RFW
|
RDW
|
RY
|
GLY
|
GLA
|
LI
|
LIQ
|
TPC
|
TFC
|
IC
|
RUT
|
ROS
|
|
PH
|
P
|
0.457**
|
0.449**
|
0.432**
|
-0.011
|
0.236*
|
0.054
|
0.112
|
0.097
|
-0.051
|
0.285**
|
0.159
|
0.156
|
|
G
|
0.854
|
0.942
|
0.936
|
0.857
|
0.379
|
0.103
|
0.202
|
0.178
|
0.008
|
0.486
|
0.189
|
0.177
|
|
PW
|
P
|
0.506**
|
0.508**
|
0.496**
|
-0.242
|
0.195*
|
0.045
|
-0.025
|
0.083
|
-0.011
|
0.156
|
0.168
|
0.076
|
|
G
|
0.894
|
0.932
|
0.954
|
0.958
|
0.396
|
0.124
|
0.075
|
0.247
|
0.112
|
0.280
|
0.236
|
0.099
|
|
MSD
|
P
|
0.417**
|
0.429**
|
0.412**
|
0.112
|
0.271**
|
0.216*
|
0.171
|
-.0173
|
0.009
|
0.189*
|
0.037
|
0.064
|
|
G
|
0.733
|
0.707
|
0.699
|
0.179
|
0.595
|
0.817
|
0.532
|
0.293
|
0.163
|
0.402
|
0.869
|
0.942
|
|
LL
|
P
|
0.142
|
0.119
|
0.132
|
0.158
|
0.192*
|
-0.059
|
0.250**
|
0.163
|
0.061
|
0.243*
|
-0.168
|
0.125
|
|
G
|
0.284
|
0.243
|
0.234
|
0.222
|
0.413
|
0.506
|
0.451
|
0.619
|
0.433
|
0.553
|
0.713
|
0.998
|
|
LW
|
P
|
0.085
|
0.117
|
0.141
|
0.176
|
0.032
|
0.155
|
0.210*
|
-0.131
|
0.021
|
-0.096
|
0.063
|
0.007
|
|
G
|
0.348
|
0.329
|
0.311
|
0.277
|
0.267
|
0.543
|
0.877
|
0.800
|
0.277
|
-0.029
|
0.243
|
0.991
|
|
NL
|
P
|
0.013
|
-0.056
|
-0.076
|
-0.089
|
0.367**
|
-0.078
|
0.310**
|
-0.873
|
0.208*
|
0.143
|
0.137
|
-0.041
|
|
G
|
0.058
|
-0.107
|
-0.115
|
-0.092
|
0.568
|
0.353
|
-0.072
|
0.656
|
0.510
|
0.404
|
0.589
|
0.971
|
|
LEL
|
P
|
0.340**
|
0.355**
|
0.381**
|
0.140
|
0.081
|
0.238*
|
0.250**
|
0.156
|
0.035
|
0.144
|
0.073
|
0.012
|
|
G
|
0.678
|
0.643
|
0.610
|
0.240
|
0.706
|
0.587
|
0.593
|
0.323
|
0.610
|
0.738
|
0.534
|
0.988
|
|
LEW
|
P
|
0.085
|
0.117
|
0.123
|
0.176
|
0.032
|
0.155
|
0.242*
|
0.053
|
0.021
|
-0.096
|
0.063
|
0.007
|
|
G
|
0.338
|
0.314
|
0.306
|
0.257
|
0.088
|
0.676
|
0.617
|
0.195
|
0.083
|
-0.187
|
0.894
|
0.953
|
|
NAB
|
P
|
0.568**
|
0.549**
|
0.571**
|
0.073
|
0.041
|
0.166
|
0.255**
|
0.050
|
-0.116
|
0.035
|
0.089
|
0.071
|
|
G
|
0.709
|
0.676
|
0.671
|
0.100
|
0.137
|
0.541
|
0.440
|
0.490
|
-0.070
|
0.105
|
0.664
|
0.919
|
|
SFW
|
P
|
0.739**
|
0.766**
|
0.782**
|
0.037
|
0.029
|
0.058
|
0.288**
|
0.050
|
-0.028
|
0.172
|
0.167
|
0.050
|
|
G
|
1.000
|
1.000
|
1.000
|
0.323
|
0.234
|
0.193
|
0.425
|
0.291
|
0-.194
|
0.385
|
0.317
|
0.953
|
|
SDW
|
P
|
0.728**
|
0.774**
|
0.784**
|
0.120
|
0.031
|
0.067
|
0.288**
|
-0.110
|
-0.042
|
0.149
|
0.075
|
0.091
|
|
G
|
1.000
|
1.000
|
0.950
|
0.491
|
0.358
|
0.045
|
0.792
|
0.341
|
0.750
|
0.510
|
0.269
|
0.949
|
|
SY
|
P
|
0.748**
|
0.763**
|
0.769**
|
0.131
|
0.051
|
0.087
|
0.297**
|
-0.105
|
-0.039
|
0.161
|
0.075
|
0.087
|
|
G
|
1.000
|
1.000
|
1.000
|
0.481
|
0.338
|
0.037
|
0.778
|
0.322
|
0.774
|
1.000
|
1.000
|
0.962
|
|
Table 4 Continued. Phenotypic (P) and genotypic (G) correlation coefficients of measured traits in G. glabra
Correlations
|
|
|
RFW
|
RDW
|
RY
|
GLY
|
GLA
|
LI
|
LIQ
|
TPC
|
TFC
|
IC
|
RU
|
ROS
|
|
RFW
|
P
|
1
|
|
|
|
|
|
|
|
|
|
|
|
|
G
|
1
|
|
|
|
|
|
|
|
|
|
|
|
|
RDW
|
P
|
0.957**
|
1
|
|
|
|
|
|
|
|
|
|
|
|
G
|
1.532
|
1
|
|
|
|
|
|
|
|
|
|
|
|
RY
|
P
|
0.989**
|
0.991**
|
1
|
|
|
|
|
|
|
|
|
|
|
G
|
1.423
|
1.342
|
1
|
|
|
|
|
|
|
|
|
|
|
GLY
|
P
|
-0.038
|
-0.019
|
-0.019
|
1
|
|
|
|
|
|
|
|
|
|
G
|
-0.198
|
-0.249
|
0.141
|
1
|
|
|
|
|
|
|
|
|
|
GLA
|
P
|
0.243*
|
0.180
|
0.180
|
0.036
|
1
|
|
|
|
|
|
|
|
|
G
|
0.453
|
0.361
|
0.360
|
0.347
|
1
|
|
|
|
|
|
|
|
|
LI
|
P
|
0.167
|
0.138
|
0.138
|
0.176
|
0.455**
|
1
|
|
|
|
|
|
|
|
G
|
0.282
|
0.350
|
0.258
|
0.894
|
0.921
|
1
|
|
|
|
|
|
|
|
LIQ
|
P
|
0.063
|
0.120
|
0.120
|
0.687**
|
-0.122
|
0.320**
|
1
|
|
|
|
|
|
|
G
|
0.313
|
0.394
|
0.289
|
0.985
|
0.597
|
0.532
|
1
|
|
|
|
|
|
|
TPC
|
P
|
-0.110
|
-0.105
|
-0.105
|
-0.276**
|
-0.336**
|
-0.276**
|
-.015
|
1
|
|
|
|
|
|
G
|
-0.264
|
-0.242
|
-0.275
|
0.689
|
0.974
|
0.159
|
1.260
|
1
|
|
|
|
|
|
TFC
|
P
|
-0.110
|
-0.129
|
-0.129
|
-0.134
|
-0.066
|
-0.132
|
-0.209*
|
0.192*
|
1
|
|
|
|
|
G
|
0.273
|
-0.401
|
-0.378
|
-0.348
|
0.444
|
-0.064
|
0.071
|
0.264
|
1
|
|
|
|
|
IC
|
P
|
0.102
|
0.078
|
0.078
|
0.109
|
0.266**
|
0.089
|
0.175
|
0.042
|
-0.016
|
1
|
|
|
|
G
|
0.292
|
0.321
|
0.217
|
0.323
|
0.831
|
0.260
|
0.674
|
0.237
|
0.262
|
1
|
|
|
|
RU
|
P
|
0.128
|
0.161
|
0.161
|
-0.263**
|
-0.040
|
0.133
|
0.177
|
-0.057
|
-0.284**
|
-0.027
|
1
|
|
|
G
|
0.344
|
0.349
|
0.294
|
0.543
|
-0.232
|
0.976
|
0.411
|
0.795
|
0.747
|
-0.184
|
1
|
|
|
ROS
|
P
|
0.056
|
0.120
|
0.120
|
-0.127
|
-0.015
|
0.153
|
-0.179
|
0.094
|
0.015
|
0.101
|
-0.209*
|
1
|
|
G
|
0.282
|
0.321
|
0.311
|
-0.354
|
-0.443
|
0.334
|
-0.291
|
0.261
|
0.192
|
0.396
|
0.432
|
1
|
|
PH: Plant height, PW: Plant width, MSD: Main stem diameter, LL: Leaf length, LW: Leaf width, NL: Number of leaflets, LEL: Leaflet length, LEW: Leaflet width, NAB: Number of axillary branches, SFW: Shoot fresh weight, SDW: Shoot dry weight, SY: Shoot yield, RFW: Root fresh weight, RDW: Root dry weight, RY: Root yield, GLY: Glycyrrhizic acid, GLA: Glabridin, LI: Liquritin, LIQ: Liquritigenin, TPC: Total phenol content, TFC: Total flavonoid content, IC: IC50, RU: Rutin, ROS: Rosmarinic acid
Table 5. Distribution of G. glabra populations in different clusters.
Cluster
|
|
Number of Populations
|
Percentage
|
populations
|
1
|
|
8
|
36.36
|
Sepidan, Taft, Bajgah, Marvast, Hajiabad, Sirjan, Bardsir, Shahrebabak,
|
2
|
|
3
|
13.64
|
Ahar , Darab, Ilam
|
3
|
|
4
|
18.81
|
Bojnord, Kashmar, Baft, Kermanshah,
|
4
|
|
7
|
31.81
|
Eqlid, Rabat, Semirom, Takestan, Kazerun, Saqqez, Yasuj
|
Table 6. Mean comparison of the measured traits on groups of cluster analysis
Character
|
|
|
|
Cluster
|
|
|
|
|
|
1
|
2
|
3
|
4
|
Mean
|
CV %
|
Plant height (cm)
|
|
66.06b
|
98.78a
|
52.55c
|
46.23c
|
60.27
|
23.27
|
Plant width (cm)
|
|
76.23a
|
87.97a
|
62.23b
|
52.40b
|
67.19
|
23.67
|
Main stem diameter (mm)
|
|
7.68b
|
9.96a
|
6.12c
|
6.10c
|
7.10
|
23.17
|
Leaf length (cm)
|
|
8.24b
|
10.71a
|
8.04b
|
7.34b
|
8.14
|
29.54
|
Leaf width (cm)
|
|
15.35a
|
16.33a
|
10.35b
|
14.35a
|
14.38
|
24.55
|
Number of leaflets
|
|
11.22b
|
13.40a
|
13.10a
|
11.11b
|
11.72
|
8.58
|
Leaflet length (mm)
|
|
27.93a
|
31.80a
|
18.42c
|
22.91b
|
24.95
|
19.78
|
Leaflet width (mm)
|
|
7.17a
|
7.60a
|
5.00b
|
6.97a
|
6.75
|
22.73
|
Number of axillary branches
|
|
5.41a
|
6.24a
|
3.66b
|
4.08b
|
4.75
|
22.80
|
Shoot fresh weight (g/plant)
|
|
338.81a
|
328.38a
|
155.44b
|
172.39b
|
251.56
|
39.86
|
Shoot dry weight (g/plant)
|
|
184.75a
|
183.50a
|
81.19b
|
98.07b
|
138.22
|
40.18
|
Shoot yield (g/m2)
|
|
577.34a
|
573.44a
|
253.71b
|
306.47b
|
431.96
|
40.18
|
Root fresh weight (g/plant)
|
|
376.16b
|
471.10a
|
223.72c
|
268.12c
|
322.73
|
31.27
|
Root dry weight (g/plant)
|
|
244.75a
|
284.34a
|
123.55b
|
165.52b
|
201.10
|
33.55
|
Root yield (g/m2)
|
|
764.85a
|
888.56a
|
386.10b
|
517.25b
|
628.25
|
33.85
|
Glycyrrhizic acid (mg/g DW)
|
|
98.33a
|
104.63a
|
67.64b
|
108.76a
|
96.64
|
37.96
|
Glabridin (mg/g DW)
|
|
0.63b
|
13.14a
|
2.41b
|
2.95b
|
2.83
|
95.82
|
Liquritin (mg/g DW)
|
|
1.62b
|
3.39a
|
0.97b
|
1.75b
|
1.70
|
67.89
|
Liquritigenin (mg/g DW)
|
|
8.18a
|
6.88a
|
4.44b
|
5.96ab
|
6.68
|
40.72
|
TPC (mg GAE/g DW)
|
|
6.31b
|
5.62b
|
7.46a
|
5.32b
|
6.14
|
21.94
|
TFC (mg QE/g DW)
|
|
16.21b
|
14.66b
|
21.00a
|
14.14b
|
16.28
|
33.42
|
IC50 (µg/ml)
|
|
79.14b
|
87.20a
|
81.51b
|
78.04b
|
79.95
|
8.27
|
Rutin (mg/g DW)
|
|
3.85a
|
3.83a
|
3.64a
|
3.63a
|
3.74
|
20.13
|
Rosmarinic acid (mg/g DW)
|
|
0.49a
|
0.56a
|
0.44a
|
0.52a
|
0.49
|
34.02
|
Table 7. Cluster analysis by construction of Mahalanobis (D2) distances between clustered groups
|
C1
|
C2
|
C3
|
C4
|
C1
|
1
|
65.63**
|
31.63**
|
12.55**
|
C2
|
|
1
|
87.17**
|
67.80**
|
C3
|
|
|
1
|
22.91**
|
C4
|
|
|
|
1
|