Abstract
The major a biotic stresses affecting chickpea production are high and low temperature, drought and salinity. Heat stress is a major a biotic stress factor, constraining chickpea production worldwide. This study was conducted to identifying chickpea genotypes combining heat tolerance and high yield potential and to study correlation among the heat tolerance indices. The genotypes were tested under different field-growing conditions, normal sown (non - heat stress) and late sown (heat stress) at two locations, Merowe and Gezira during winter season 2018/2019. The trials were laid out in alpha lattice design with three replications. Eight heat tolerance indices, which were most frequently used in plant breeding including, geometric mean productivity (GMP), yield index (YI), mean productivity (MP), stress susceptibility index (SSI), stress tolerance index (STI), tolerance index (TOL), sensitivity heat index (SHI) and relative heat index (RHI) were calculated based on seed yield under heat stress (Ys) and non - heat stress (Yp) conditions. Under both environments, the results from combined analysis of variance showed that there were highly significant differences among the genotypes for all traits studies. There were highly significant differences observed among the tested chickpea genotypes response to heat stress. Seed yield in stress (Ys) condition was positive and highly significant correlated with geometric mean productivity, yield index, stress tolerance index and relative heat index. The Ys was negative and highly significant correlated with stress susceptibility index and sensitivity heat index. Seed yield in non-stress (Yp) condition was positive and significant correlated with geometric mean productivity, yield index, stress tolerance index and tolerance index. The results of this research showed that the six indices, GMP, YI, SSI, STI, SHI and RHI can be used as optimal indicators for screening heat tolerant chickpea genotypes. Nine genotypes including four released improved varieties (Shiekh Mohamed, Wad Hamid, Salwa and Hwata) and five genotypes (no. 11, 4, 26, 30 and 34) are most stable chickpea genotypes for heat tolerance and can be further used in breeding program. These genotypes can be used in the chickpea national breeding program to develop varieties with improved heat tolerance in Sudan. Genotype no. 11 (FLIP 08-59 C) was release by Sudan national variety release committee for commercial production under the name Elbarkal in September 2022 to will be grown in Gezira, River Nile and Northern States of Sudan.
Author Contributions
Copyright© 2023
E. A. Hamza Fatih, et al.
License
This work is licensed under a Creative Commons Attribution 4.0 International License.
This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Competing interests The authors have declared that no competing interests exist.
Funding Interests:
Citation:
Introduction
Among the food legumes, chickpea occupies second position after beans. It is cultivated in more than fifty countries worldwide In Sudan, chickpea is the third most economically important food legume crop after faba bean and cowpea, as a cash crop that generates income for farmers and rural communities, and as a significant source of protein for Sudanese people Chickpea reproductive stages (flowering and podding) are vulnerable to external environmental changes and heat stress Several indices based on the yield under non-heat stress (Yp) and heat stress (Ys) have been introduced for the selection of heat tolerant genotypes. Among these, the indices employed in various stress conditions are stress tolerance (TOL) and mean productivity (MP) introduced by some researchers To date, limited genetic resources for heat stress tolerance in chickpea have been reported Thus, achieving optimum grain yield underheat-tolerant heat stress remains prime criteria to chickpea breeders. Implication of various indices helps in measuring yield loss and screening heat stress tolerant genotype under stress condition
Materials And Methods
The study area of this research was conducted over one consecutive winter season 2018/2019 in two locations in the Agricultural Research Corporation (ARC) of Sudan. Namely the Gezira Research Station Farm (GRSF), located in central clay plain of Sudan within the latitude 14°24' N, and longitude 33°29' E and altitude 407 meters above sea level with soil characterized by cracking heavy clay (vertisols), very low water permeability, pH of 8.3, organic matter (0.4%), nitrogen (0.04% ppm), and phosphorus (ESP, 4 ppm) and (2) farmers field in the Northern state of Sudan, Merowe locality ( latitude: 18° 27' 0" N, longitude: 31° 49' 59" E, elevation: 258 meters above sea level). Forty three chickpea genotypes were selected from advanced materials of the national chickpea breeding program. In addition, five commercial chickpea cultivars which were released by the Agricultural Research Corporation of Sudan namely, Shiekh Mohamed, Merowe, Wad Hamid, Salwa and Hwata ( Across all growing locations, the land was prepared by disc ploughing, disc- harrowing, leveling and ridging. In two locations the genotypes were evaluated in two environments i.e., normal sown (15 November) and late sown (5 December). The experiment was conducted in Alpha Lattice design with three replications under normal sown and late sown conditions. Each replicate consisted of twelve incomplete blocks and four plots in each block. Each genotype was planted in a separate plotwhich was consisted of one ridge /row of 4 meter length, witha plant-to-plant and row-to-row distance of 10 cm and 60 cm,respectively. The plots were separated by a distance of 60cm. Irrigation was carried out at 12-14 days intervals to avoid any water stress. A starter dose of nitrogen in the form of urea urea was applied at a rate of 43 kg N/ha with the third irrigation. Weeds were kept to the minimum by hand removal during the first month after sowing. Seed yield was assessed from a net area of 2.4 m2 (one row x 4 m long x 0.6m). (a) Days to 50% flowering: Days to 50% flower initiation was recorded as the number of days required from planting to the time when 50% of plants in plots produced at least one flower. (b) Days to 90% physiological maturity: Days to 90% physiological maturity was recorded as the number of days required from planting to the time when 90% of plants showed a yellow color in each plot before senescence. (c) Plant height: Plant height was recorded from ten randomly taken plants from one central row at physiological maturity from ground to the tip of the main stem and then the mean was recorded as height per plant (cm). (a) Number of pods per plant: It was counted from the total number of seed bearing mature pods was counted separately from randomly selected five plants in each plot and averaged. (b) Number of seeds per plant: It was counted from the sample after threshing, as counted from each five randomly taken plants per middle of row and then expressed as an average of 5 plants. (c) Number of seeds per pod: This was calculated by dividing the total number of seeds per plant (of the sample) by the total number of pods per plant. (d) Hundred-seed weight (g): It was calculated by 100 seed samples were randomly selected from each plot and seeds weighted. (e ) Seed yield per plant (g): It was calculated as the total seed produced from five randomly selected plants after threshing and cleaning was weighted in gram with the help of electronic top pan balance and averaged out for seed yield per plant (g). Harvest index (HI): It was calculated as the ratio of economic yield divided to the total of biological yield expressed in percentage. HI (%) = (Seed yield / Biological yield) x 100 (f) Biomass (t ha-1): Biological yield is the total yield of crop including economic yield and the straw yield. The biological yield was recorded after harvesting using electronic balance (in g or kg net plot) and converted into (t ha-1). (g) Seed yield (t ha-1): When signs of maturity were clear on the plant (complete yellowing of leaves), four meter length in each plot was harvested for yield, weighed and then seed yield per plot was converted to seed yield in ((t ha-1) adjusted at 10% moisture content. Eight heat tolerance indices namely, geometric mean productivity (GMP), yield index (YI), mean productivity (MP), stress susceptibility index (SSI), stress tolerance index (STI), tolerance index (TOL), sensitivity heat index (SHI) and relative heat index (RHI) were calculated by the given formulae:- Geometric mean productivity (GMP) Yield index (YI) Mean productivity (MP) = (Ypi+Ysi) / 2. The genotypes with high values of this index found to be more desirable Stress susceptibility index (SSI) = (1-(Ysi/Ypi))/SI. The genotypes with least SSI values have tolerant to heat stress Stress tolerance index (STI) = (Ypi/Yp) (Ysi/Ys) (Ys/Yp) The genotypes with high STI values have tolerant to heat stress Tolerance index (TOL) Sensitivity heat index (SHI) = (Ypi - Ysi) / Ypi. The genotypes with least SHI values have tolerant to heat stress Relative heat index (RHI) = (Ysi / Ypi) / (Ys/Yp). The genotypes with maximum RHI values have tolerant to heat stress % Reduction = (Ypi-Ysi)/Ypi x 100 In the above formulas Ysi, Ypi, Ys and Yp represent yield under heat stress, yield under non-heat stress for each genotype, yield mean in heat stress and non-heat stress conditions for all genotypes, respectively. The collected data were subjected to combine analysis of variance using the GenStat 12th edition statistical analysis package for windows (2009) to test the level of significance among the genotypes for different traits under non - heat stress and heat stress conditions. The correlation analysis was worked out according to the method described by
FLIP 09 - 181 C
X06TH53/FLIP03-128CXFLIP01-25C
ICARDA
LIP 09 - 179 C
X06TH53/FLIP03-128CXFLIP01-25C
ICARDA
FLIP 09 - 184 C
X06TH53/FLIP03-128CXFLIP01-25C
ICARDA
FLIP09 - 155 C
X06TH71X05TH104XFLIP03-39C
ICARDA
FLIP09 - 438 C
S00794(60 KR)-44
ICARDA
FLIP09 - 261 C
X04TH32/X03TH-32XFLIP99-34
ICARDA
FLIP 07 - 236 C
X03TH138/FLIP98-130CXFLIP99-34C
ICARDA
FLIP 09 - 259 C
X04TH32/X03TH-32XFLIP99-34C
ICARDA
FLIP08 - 86 C
X03TH144/FLIP97-116CXFLIP97-32C
ICARDA
FLIP09 - 6 C
X05TH22/(FLIP99-46CXFLIP97-91C)XFLIP02-43C
ICARDA
FLIP 08-59 C
X02TH3/FLIP 98-28C X FLIP-97-102C
ICARDA
FLIP 09-182 C
X06TH53/FLIP03-128C X FLIP01-25C
ICARDA
FLIP 09-187 C
X06TH53/FLIP03-128CXFLIP01-25C
ICARDA
FLIP09 - 240 C
F2X01TH186 (45KR)-3
ICARDA
22330
70755
ICARDA
22304
70381
ICARDA
22317
70309
ICARDA
22233
9440
ICARDA
22278
70334
ICARDA
22267
70304
ICARDA
22232
9439
ICARDA
22223
9425
ICARDA
22235
9442
ICARDA
22366
74021
ICARDA
22293
70357
ICARDA
22380
1.00E+05
ICARDA
22362
73386
ICARDA
22254
70273
ICARDA
22335
70764
ICARDA
22204
6057
ICARDA
22272
70312
ICARDA
222389
1.00E+05
ICARDA
222303
70379
ICARDA
222242
69620
ICARDA
22373
75360
ICARDA
22206
6109
ICARDA
22384
70780
ICARDA
22341
70773
ICARDA
22302
70377
ICARDA
22260
70286
ICARDA
22266
70299
ICARDA
22392
1.00E+05
ICARDA
22261
70290
ICARDA
Shiekh Mohamed
X99TH62/(FLIP932CxFLIP 94-115C)
Released commercial cultivar
Merowe
X99TH62/(FLIP932CxFLIP 94-115C)
Released commercial cultivar
Wad Hamid
(India-ICRISAT Selection)
Released commercial cultivar
Salwa
(X87TH 186/ ICCI 4198)//FLIP 82-150C )
Released commercial cultivar
Hwata
(ICCV2/Surutato 77)//ICC 7344 )
Released commercial cultivar
Discussion
The results of the analysis of variance for different traits under non-heat stress and heat stress conditions are presented in DF: Number of days to 50% flowering, DM: Number of days to 90% maturity, PH: Plant height (cm), NPP: Number of pods per plant, NSPL: Number of seeds per plant, NSP: Number of seeds per pod, 100-SW: Hundred seed weight, SYP: Seed yield per plant (g), HI (%): Harvest index, BIO: Biomass (t ha-1) and SY: Seed yield (t ha-1). *, ** and *** Significant at 0.05, 0.01 and 0.001 probability levels, respectively. n.s = non - significant difference at 0.05 probability level. Similarly, previous studies on chickpea landraces also reported by Different heat tolerance indices were calculated on the basis of seed yield of the chickpea genotypes under normal sowing date (non-heat stress) (Ypi) and late sowing date (heat stress) (Ysi) conditions are presented in ( GMP = Geometric mean productivity, YI = Yield index, MP = Mean productivity, SSI = Stress susceptibility index, STI = Stress tolerance index, TOL = Tolerance index, SHI = Sensitivity heat index and RHI = Relative heat index. Comparison of genotypes based on reduction of yield and yield related traits by growing them under non-heat stress and heat stress environments could be an important parameter for selection of tolerant genotype under drought, heat and other a biotic stresses in crop plants The genotype no. 11 and the check Wad Hamid (no. 46) exhibited excellent seed yield in heat stress (2.52 and 2.35 t ha- Considering YI as an important selection index for heat tolerance, genotypes no. 11(1.65), 47 (1.57), 48 (1.49), 4 (1.39), 44 (1.36), 43 (1.30), 34 (1.27), 30 (1.25) and 26 (1.22), showed higher YI value than the check Merowe (1.15) ( Thus genotypes exhibiting higher MP, GMP and YI could be efficiently used in selecting superior genotypes under heat stress. While emphasizing SSI as another important selection parameter for heat tolerance, genotypes showing SSI<1 were considered as higher heat tolerance Study To determine the most desirable heat tolerant criteria, the correlation coefficients between Yp, Ys and other quantitative indices of heat tolerance were calculated ( GMP: Geometric mean productivity, (YI): Yield index, (MP): Mean productivity, (SSI): Stress susceptibility index, (STI) , Stress tolerance index, (TOL): Tolerance index, (SHI): Sensitivity heat index, (RHI): Relative heat index, Ypi: Mean seed yield of individual genotype in non - heat stress condition and Ysi: Mean seed yield of individual genotype in heat stress condition. ** and *** Significant at 0.01 and 0.001 probability levels, respectively. n.s = non - significant difference at 0.05 probability level. It was found that seed yield under heat stress condition (Ys) showed negative and highly significant correlation with SSI Considering MP, it had high and positive correlation with GMP, YI, STI, RHI and Yp but it had negative correlation with TOL and SHI. STI showed high and positive correlation with GMP, YI, RHI, Yp and Ys, but it indicated negative association with SSI and RHI. Both SSI and TOL showed high significant negative association with RHI, whereas SSI and TOL exhibited high positive correlation.
Traits
Location (L)
Genotype (G)
L x G
Non-heat stress
Heat stress
Non-heat stress
Heat stress
Non-heat stress
Heat stress
DF
1200.50***
2.00n.s
364.92***
295.72***
36.02***
55.48n.s
DM
20334.72***
11312.59***
156.04***
80.88***
126.49***
64.37***
PH
4985.01***
8109.01***
142.92***
120.37***
33.17n.s
61.11***
NPP
161420.9***
82872.0***
2254.4***
1166.2***
1812.4***
1111.3***
NSPL
303031.1***
147451.7***
3962.7***
2006.9***
2412.2***
1678.2***
NSP
1.87857***
1.77818***
0.12627***
0.12043***
0.02964n.s
0.03048n.s
100-SW
703.12***
1984.50***
246.21***
268.15***
27.20**
19.49*
SYP
32340.60***
24708.65***
223.21***
198.02***
179.08***
143.18***
HI (%)
5352.68***
1880.38***
122.50***
91.64***
125.81***
97.10***
BIO
669911156***
826400961***
10556848***
7051932***
9667824***
5703943***
SY
1439480n.s
61633678***
1929551**
1029497***
1115749n.s
551317***
3.93
1.52
61.4
2442
0.99
2725
1.26
0.64
2416
0.61
0.75
2.61
1.36
47.9
1880
0.89
1981
0.98
2.37
1251
0.47
1.01
3.78
1.32
65
2233
0.86
2549
1.33
2.65
2457
0.65
0.68
3.45
2.12
38.6
2704
1.39
2786
0.79
3.07
1336
0.38
1.19
3.21
1.75
45.3
2370
1.15
2479
0.93
2.74
1455
0.45
1.06
3.64
1.3
64.2
2175
0.85
2470
1.32
2.59
2338
0.64
0.69
2.74
1.37
49.8
1941
0.9
2058
1.02
2.34
1368
0.5
0.97
2.72
1.05
61.4
1690
0.68
1886
1.26
2.12
1674
0.61
0.75
2.82
1.05
62.6
1721
0.69
1935
1.28
2.15
1766
0.62
0.72
3.45
1.42
58.7
2214
0.93
2436
1.2
2.61
2028
0.58
0.8
3.29
2.52
23.3
2881
1.65
2907
0.48
3.27
770
0.23
1.49
2.58
1.74
32.4
2119
1.14
2161
0.66
2.52
838
0.32
1.31
3.2
1.77
44.7
2381
1.16
2487
0.91
2.75
1434
0.44
1.07
3.51
1.61
54.1
2376
1.05
2559
1.11
2.75
1899
0.54
0.89
2.83
1.29
54.5
1909
0.84
2059
1.12
2.31
1545
0.54
0.88
2.84
1.63
42.5
2154
1.07
22238
0.87
2.54
1210
0.42
1.11
2.2
1.09
50.4
1550
0.71
1647
1.03
1.97
1112,
0.5
0.96
3
1.5
49.9
2121
0.98
2250
1.02
2.51
1498
0.5
0.97
3.45
1.7
50.8
2418
1.11
2573
1.04
2.79
1756
0.5
0.96
2.18
1.12
48.5
1566
0.73
1653
0.99
1.98
1059
0.48
1
3.16
1.42
54.8
2120
0.93
2290
1.12
2.51
1733
0.55
0.87
2.55
1.47
42.1
1937
0.96
2010
0.86
2.34
1075
0.42
1.12
3.04
1.12
63.2
1845
0.73
2081
1.29
2.27
1925
0.63
0.71
2.88
1.14
60.3
1816
0.75
2014
1.24
2.23
1740
0.6
0.77
2.9
1.44
50.3
2042
0.94
2168
1.03
2.43
1459
0.5
0.96
2.73
1.86
31.9
2250
1.22
2292
0.65
2.65
872
0.31
1.32
3.4
1.69
50.2
2399
1.11
2546
1.05
2.77
1707
0.5
0.96
2.09
0.83
60
1318
0.54
1459
1.23
1.76
1251
0.6
0.77
2.94
1.5
48.8
2100
0.98
2219
1
2.49
1435
0.48
0.99
3.14
1.91
39.1
2447
1.25
2524
0.8
2.82
2130
0.39
1.18
2.4
1.51
37.1
1900
0.99
1952
0.76
2.31
891
0.37
1.22
2.87
1.1
61.6
1773
0.72
1982
1.26
2.2
1768
0.61
0.74
1.86
0.98
47.2
1353
0.64
1423
0.97
1.78
880
0.47
1.02
3.22
1.94
39.5
2500
1.27
2581
0.81
2.87
1272
0.39
1.17
2.74
1.44
47.3
1986
0.94
2089
0.97
2.38
1295
0.47
1.02
2.2
1.61
26.7
1885
1.06
1908
0.55
2.31
589
0.26
1.42
2.03
0.89
56.2
1341
0.58
1458
1.15
1.78
1142
0.56
0.85
3.26
1.16
64.3
1945
0.76
2211
1.32
2.37
2100
0.64
0.69
3.59
1.56
56.6
2361
1.02
2571
1.16
2.74
2032
0.57
0.84
3.93
1.31
66.7
2264
0.85
2616
1.37
2.69
2620
0.66
0.64
1.84
0.93
49.3
1307
0.61
1384
1.01
1.74
907
0.49
0.98
1.91
1.07
44
1431
0.7
1492
0.9
1.86
844
0.43
1.09
3.58
1.99
44.4
2670
1.3
2787
0.91
3.03
1594
0.44
1.08
3.02
2.08
31
2509
1.36
2553
0.63
2.9
940
0.31
1.34
3.91
1.76
54.8
2624
1.15
2835
1.12
2.99
2144
0.54
0.87
2.58
2.35
8.9
2459
1.54
2462
0.18
2.92
232
0.08
1.77
3.51
2.4
31.4
2900
1.57
2953
0.64
3.27
1104
0.31
1.33
3.53
2.28
35.3
2834
1.49
2902
0.72
3.2
1247
0.35
1.25
2.96
1.52
0.9042***
0.1587
0.1579
-0.3857**
-0.7383***
-0.0967
0.7554***
0.7578***
0.1257
-0.4333**
0.1839
-0.2390
-0.0173
0.7942***
-0.0283
-0.3859**
-0.7380***
-0.0977
0.9996***
-0.4324**
0.7937***
0.3870**
0.7391***
0.0944
-0.9998***
0.4322**
-0.7931***
-0.9997***
0.8046***
0.4770***
0.0995
0.2235
0.5032***
0.7118***
0.2224
-0.2219
0.9058***
0.9999***
0.1563
-0.7357***
0.7577***
-0.2355
-0.7353***
0.7366***