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Genetic Variation Studies of Ionic and within Boll Yield Components
in Cotton (Gossypium Hirsutum L.) Under Salt Stress
Muhammad Mubashar Zafar
a,b
*, Abdul Razzaq
a,c
*, Muhammad Awais Farooq
b
*,
Abdul Rehman
a,b,d
, Hina Firdous
d
, Amir Shakeel
b
, Huijuan Mo
a,e
, Maozhi Ren
a,e
,
Muhammad Ashraf
f
, and Yuan Youlu
a
a
Institute of Cotton Research, Chinese Academy of Agricultural Sciences, Anyang, China;
b
Department of Plant
Breeding and Genetics, University of Agriculture Faisalabad, Faisalabad, Pakistan;
c
Institute of Molecular Biology and
Biotechnology, the University of Lahore, Lahore, Pakistan;
d
Department of Plant Pathology, University of Agriculture
Faisalabad, Faisalabad, Pakistan;
e
Zhengzhou Research Base, State Key Laboratory of Cotton Biology, Zhengzhou
University, Zhengzhou, China;
f
University of Agriculture Faisalabad, Faisalabad, Pakistan
ABSTRACT
Cotton (Gossypium hirsutum L.) productivity is decreasing alarmingly by
salinity. For genetic analysis of cotton response to salinity 8 parental geno-
types and their 16 F
1
crosses were evaluated in Line × Tester fashion under
normal and salt stress 15 dSm
−1
. Data were recorded for plant height,
number of bolls plant
−1
, boll weight, lint weight, seed cotton yield plant
−1
,
lint percentage, seed index, lint index, number of seeds boll
−1
, seed mass
boll
−1
, lint mass boll
−1
, lint mass per seed, seed volume 100
−1
seeds, seed
surface area, ionic ratio, ber strength, ber length, ber neness, GOT %,
SOD, POD, CAT, TSP, and H
2
O
2
. The results showed that SCA estimates were
higher than the GCA indicating the preponderance of non-additive gene
action. Kahkashan, MS-71, and CRS-2007 were found good general combiner
whereas FH-114 × FH-312 was found the best specic combiner. FH-114
× KAHKASHAN and FH-114 × FH-312 demonstrated signicant heterosis for
most traits under normal and salt stress conditions. The present study
revealed that salt tolerance ability is controlled by a few dominant genes in
the identied genotypes.
摘要
棉花（Gossypium hirsutum L.的生产力正因盐度而急剧下降. 对16个杂交组
合进行了盐胁迫下的亲本基因型分析. 记录了株高株铃数1铃重皮棉产量株
1皮棉百分率种子指数皮棉指数籽铃数1种子质量铃1皮棉质量1每粒皮棉
质量种子体积100-1粒种子比表面积离子比纤维强度纤维长度纤维细度
GOT%SODPODCATTSP和H2O2. 结果表明SCA估计值高于GCA表明非加性基
因作用占优势. KahkashanMS-71和CRS-2007被认为是良好的通用合路器而
FH-114×FH-312是最佳的专用合路器. 在正常和盐胁迫条件下FH-
114×KAHKASHAN和FH-114×FH-312多数性状表现出显著的杂种优势. 本研
究表明在已鉴定的基因型中耐盐能力受少数显性基因控制.
KEYWORDS
Cotton; salinity; Line ×
Tester; gene action; ionic
homeostasis; fiber traits;
antioxidants; hydrogen
peroxide
关键词
棉花; 盐度; 线路×测试仪;
基因作用; 离子稳态; 纤维
特性; 抗氧化剂; 过氧化氢
Introduction
Cotton plays an important role in the economy of Pakistan with the contribution of 0.8% to GDP and
has an agriculture value addition of 4.5%. Its production in 2018–19 was 9.861 million bales that
CONTACT Abdul Razzaq dramirpbg@gmail.com Institute of Cotton Research, Chinese Academy of Agricultural Sciences,
Anyang 455000.
*These authors contributed equally to this work.
Abbreviations: H
2
O
2
= Hydrogen peroxide; SOD= Superoxide dismutase, POD= peroxidase; CAT= Catalase; TSP= Total soluble
proteins; GOT= ginning out turn
JOURNAL OF NATURAL FIBERS
https://doi.org/10.1080/15440478.2020.1838996
© 2020 Taylor & Francis
indicated a decline of 17.5% in comparison to the previous year (Anonymous 2018-19). This
significant decline in cotton production is owing to various environmental factors (Ismail and
Horie 2017) and salt stress is found to be the most severe environmental factor that has rendered
40% of the world area as sodic (Farooq et al. 2017). Salinity has been increased in an agricultural area
with a drastic change in climate, i.e., unpredictable temperature fluctuation and rainfall pattern
(Arzani 2008), unscientific irrigations as well as excessive use of fertilizers (Munns and Tester
2008). Low cotton yield and high population pressure have increased the demand for cotton (Sattar,
Hussnain, and Javaid 2010). It is very important to enhance its production in cultivated areas of
Pakistan in order to meet population pressure.
Cotton has moderate tolerance against salinity (Du et al. 2016), it can tolerate up to 7.7 dSm
−1
(Wei
et al. 2017). The osmotic stress, ionic toxicity (Flowers and Colmer 2008), essential nutrient deficiency
(Ahmad et al. 2013), reduced photosynthetic rates (Abogadallah 2010), and abnormal physiological
processes in plants are associated with salinity stress (Hameed et al. 2014). A higher level of soil salinity
causes a reduction in seedling emergence, decrease vegetative and reproductive growth leading to low
cotton yield and poor fiber quality (Dong 2012). The concentration of sodium ions outside the cell
leads to sodium ions accumulation inside the cell due to the concentration gradient. The excess of
sodium ions leads to the toxicity inside the cell that halts the metabolic process. It also damages the
root cells and renders the cell inefficient to expel the sodium ions outside to generate a homeostatic
environment (Farooq 2019). The ionic toxicity leads to the production of poor quality of cotton fibers
and low yield. Therefore, the impact of salinity on the cotton can be better understood by employing
modern biotechnological, biochemical, molecular markers, and conventional techniques (Nabi et al.
2011). The use of biochemical markers to identify the salt-tolerant genotypes has become very reliable.
The induction of salt stress kick starts the production of reactive oxygen species (ROS), i.e., hydrogen
peroxide that are battled down by the production of antioxidants. The quantity of antioxidants, i.e.,,
super-oxidase dismutase, peroxidases, catalase, total soluble proteins can dictate the ability of the plant
to withstand salt stress (Ma, Dong, and Li 2011).
Cotton has natural variation against salt stress that can be utilized in the screening and develop-
ment of salt-tolerant cotton genotypes. Earlier studies have reported the impact of salt stress in cotton
crops at the germination and seedlings stage (Farooq et al. 2019) whereas, the detailed analysis and
interaction of agronomic, fiber, and biochemical traits have not been met yet. The present study will
not only study the impact of high salt stress on the cotton crop but also assist in future breeding
programs by providing salt-tolerant germplasm.
Materials and methods
The research was conducted in the experimental area of the Department of Plant Breeding and
Genetics, University of Agriculture Faisalabad. The seeds of eight cotton varieties were sown in
earthen pots in glass house (Table 1). The crosses were made in line × tester fashion to get F
1
seed. For this purpose, four genotypes, namely (MS-71, FH-114, IUB-65 NS-131) were used as
a line and four genotypes (Kahkashan, CRS-2007, FH-312, CIM-573) as a tester. The optimal
growing conditions, i.e., temperature, humidity, and rain fall are given in Figure 1, Table 2. After
crosses, selfed and crossed bolls were collected separately at the time of boll opening. After
picking their ginning was done separately by a single roller ginning machine (Testex, Model:
TB510C). Their seeds were preserved in butter paper bag to protect them from contamination
Table 1. The material used in the breeding study.
Lines Testers Crosses Crosses Crosses Crosses
MS-71 Kahkashan MS-71× Kahkashan FH-114× Kahkashan IUB-65× Kahkashan NS-131× Kahkashan
FH-114 CRS-2007 MS-71× CRS-2007 FH-114× CRS-2007 IUB-65× CRS-2007 NS-131× CRS-2007
IUB-65 FH-312 MS-71× FH-312 FH-114× FH-312 IUB-65× FH-312 NS-131× FH-312
NS-131 CIM-573 MS-71× CIM-573 FH-114× CIM-573 IUB-65× CIM-573 NS-131× CIM-573
2M. M. ZAFAR ET AL.
till the time of the next sowing of the crop. In the next season the seeds of 24 genotypes
containing 8 parents and 16 crosses were sown in earthen pots with three replications by using
split-plot design under randomized complete block design RCBD. The seeds of all genotypes
were sown in controlled (without salt) and salt mixed earthen pots having a salinity of 15 dsm
−1
.
These pots were kept in an open field from sowing to harvesting of the crop. The level of salinity
(15 dSm
−1
) was maintained in earthen pots by using US Regional Laboratory (1954) method. To
maintain the required salt concentration the EC (soil electrical conductivity) of pots was
examined after every two days and was maintained during the experiment. All standard agro-
nomic practices were performed properly from sowing to harvesting. At maturity (> 80% boll
formation completed) the data were taken on the following traits; Plant height, number of bolls
plant
−1
, Boll weight, lint weight, Seed cotton yield plant
−1
, Lint percentage, Seed index, Lint
index, No. of seeds boll
−1
, Seed mass boll
−1
, Lint mass boll
−1
, Lint mass per seed, Seed volume
per 100 seeds, seed surface area, fiber strength, fiber length, fiber fineness, ginning out turn
percentage (GOT%), Na
+
, K
+
, K
+
/Na
+
ratio, SOD, POD, CAT, TSP, and hydrogen peroxide from
five selected plants from each replication of both controlled and saline treatment. The recorded
data were subjected to analysis of variance (Steel, Torrie, and Dickey 1997). Line × tester analysis
of (Kempthorne 1957) was used for the estimation of combining ability effects of parental
genotypes and crosses. Heterosis effects were calculated by the method outlined by Mather
and Jinks. (Mather and Jinks 2013).
Table 2. Metrological data during the cotton growing season.
Months
Temperature (
o
C) Relative humidity (%) Rain fall (mm)
Max. Min. Avg. Avg. Avg.
June 47 22 33.9 56.5 92
July 44 21 33 70.2 195.8
August 41.5 26 34.2 67.4 5.4
September 43 21 31.4 65.1 41.7
October 36 15.5 25.6 64.4 0
November 30.5 8.5 19.7 74.6 0.6
December 27 1.5 14.1 81.5 0.7
Figure 1. Optimal climatic conditions for cotton growth and development.
JOURNAL OF NATURAL FIBERS 3
Fiber quality traits
A representative sample from seed cotton was weighed and ginning was done by a single roller ginning
machine. The seeds were separated from the lint and ginning out turn was calculated by dividing the
weight of lint in a sample by seed cotton weight of the sample, which was expressed in percentage. Lint
was further processed to take out the parameters of fiber fineness, fiber strength, and fiber length with
high volume instrument (HVI-900, USTER, USA).
Ionic analysis
The sodium and potassium analysis were calculated by taking fresh green leaves at noon, when the
plants had reached their vegetative maturity stage and then they were dried in hot air. Dried leaves
were ground down in mortar and pestle and then the leaves were digested with nitric acid and sulfuric
acid (2:1 molar ratio) on the hot plate. When the digestion completed, the material was cooled down to
room temperature and readings were taken by a flame photometer (410 Flame Photometer). The
potassium to sodium ratio was estimated by dividing potassium concentration to sodium
concentration.
Hydrogen peroxide assay
H
2
O
2
was measured by following the method of Bernt and Bergmeyer (Allen, Farmer, and Sohal
1983). The H
2
O
2
was quantified by taking 0.5 g of leaf samples from control and treatment groups
were homogenized with liquid nitrogen, and the powders were suspended in 1.5 mL of 100 mM
potassium phosphate buffer (pH 6.8). Suspensions were then centrifuged (Refrigerated SIGMA 2–
16KL Centrifuge, UK) at 18,000 × g for 20 min at 40°C. The enzymatic reaction was initiated taking
0.25 mL supernatant and 1.25 mL peroxidase reagent consisting of 83 mM potassium phosphate buffer
(pH 7.0), 0.005% (w/v) O-dianizidine, 40 μg peroxidase/mL at 30°C. The reaction was stopped after
10 min by adding 0.25 mL of 1 N perchloric acid and the reaction mixture was centrifuged at 5000 × g
for 5 min. The absorbance was checked at 436 nm through spectrophotometer (NanoDrop™ 8000
Spectrophotometer Thermo Fisher Scientific, Sweden), and the amount of H
2
O
2
was calculated using
an extinction coefficient of 39.4 mM
−1
cm
−1
(Zhang et al. 2014).
SOD assay
Super-oxidase dismutase was quantified as units of enzymes that inhibited the photochemical
reduction of nitro blue tetrazolium (NBT). The reaction mixture was consisted of potassium
phosphate buffer (pH 5) + 200 μL methionine + 200 μL Triton X + 100 μL NBT + 800 μL distilled
water dissolved with enzyme extracts of 100 μL and were placed under ultraviolet light for
15 minutes and then 100 μL of Riboflavin was added. The readings were taken on a spectro-
photometer at 560 nm absorbance.
POD assay
Peroxidase (POD) values were calculated as the amount of enzymes unit that oxidized guaiacol.
Enzyme extracted for SOD was used for measuring POD activity. The reaction mixture consisted of
800 μL potassium phosphate buffer (pH 5) + 100 μL H
2
O
2
(40 mM) + 100 μL guaiacol (20 mM) was
mixed with the 100 μL of enzyme extract in the Eppendroff tubes and readings were taken at 470 nm
wavelength through spectrophotometer (Liu et al. 2009).
4M. M. ZAFAR ET AL.
Catalase assay
The catalase contents were observed as the amount of H
2
O
2
consumed by catalase and converted into
H
2
O and O
2
. Same enzyme extract of 100 μL quantity was used that was prepared for SOD and was
mixed with 100 μL of H
2
O
2
in the cuvettes, and readings were taken at 240 nm absorbance through
spectrophotometer (Liu et al. 2009).
Total soluble proteins
The leaf tissues were extracted for the measurement by using potassium phosphate buffer (pH 4),
vortexed, and centrifuged. The same enzyme extract was taken in 40 μL quantity that was prepared for
SOD and was mixed with 160 μL quantity of Bradford reagent. The mixture was added into the ELISA
plate and readings were recorded through spectrophotometer at 595 nm absorbance (Bradford 1976).
Results
Analysis of variance for various genotypes of G. hirsutum under normal and saline
conditions in Line × Tester design
Analysis of variance following line × tester technique was carried out to assess to presence of genetic variations
among plant material grown in normal and saline conditions (Tables 3 & 4). Biometrical analysis revealed that 16
crosses along with their 8 parents differed significantly from each other, however, lint percentage, seed per boll
and Na
+
indicated non-significant differences under normal condition, moreover, under saline conditions boll
weight, lint percentage lint index, seed per boll and lint mass per seed exhibited non-significant differences.
Under normal conditions crosses showed significant differences in all traits except lint percentage, seed per boll,
Na
+
and hydrogen peroxide whereas under salt stress boll weight, lint percentage, seed per boll, lint mass per seed,
seed volume, seed surface area and super-oxidase dismutase were not significantly different from each other.
Lines × Testers showed significant differences for all traits under normal conditions except lint percentage, seed
per boll, lint mass per boll, lint mass per seed, K
+
/Na
+
, Na
+
and hydrogen peroxide. Crosses vs parents showed
significant differences for lint percentage, seed index, seed mass per boll, fiber length, K
+
, hydrogen peroxide and
super-oxidase dismutase under normal and salt stress conditions.
General Combining ability eects under normal and saline conditions
Comparison of estimates for plant height reveals that line FH-114 exhibited good general combiner in
normal (−7.12) and saline (−5.44) conditions than the rest of the all. Whilst testers CRS-2007, CIM-
573 showed significant GCA effects under both normal (2.88 & 9.87) and saline (1.91 & 6.24)
conditions (Tables 5 and 6). For the number of bolls per plant, the lines IUB-65 (1.19) showed positive
and significant GCA effects under normal condition while MS-71 (0.85) under saline conditions.
Among the testers the CRS-2007 (1.69, 1.52) exhibited good general combining ability under normal
and saline conditions. For boll weight, all lines showed non-significant GCA effects under normal and
saline conditions. Tester, CRS-2007 showed good general combining ability under both conditions
(0.31, 0.36). Line MS-71 showed positive and significant estimates (2.63) for seed cotton yield under
normal conditions whereas Tester CRS-2007 was found good general combiner under saline condi-
tions. For lint mass per boll, line IUB-65 showed significant GCA results whereas tester CRS-2007
displayed positive and significant GCA under both conditions for lint mass per boll and lint mass
per seed. Estimates for seed surface revealed that IUB-65 and MS-71 were good general combiner
under normal and saline conditions, respectively. For fiber fineness and fiber length, line MS-71 and
tester CRS-2007 indicated positive and significant results under normal and salt stress. Whereas
Kahkashan showed the highest GCA (1.73) for GOT under normal conditions. All the lines had non-
significant GCA for K
+
/Na
+
ratio in both conditions except NS-131 in normal condition whereas the
tester FH-312 exhibited positive and significant GCA (0.44) under saline condition. For Na
+
conc.
JOURNAL OF NATURAL FIBERS 5
Table 3. Mean square values of line × tester analysis for different traits under normal and saline conditions.
SOV DF Trt PH NB BW LW SCY L% SI LI S/B SM/B LM/B LM/S SV SSA
Rep. 2 N 4.9164 9.4306** 0.1454 17.6732** 98.6717** 73.7293 2.418** 2.3289 5.4766 4.9444* 0.9448* 0.0023* 99.0139 0.0002**
S 4.7135 6.4306* 0.0372 4.4714 26.1897* 9.9207 0.5383 0.2886 11.2789 3.5250* 0.5940* 0.0007 1.6250 0.0002
Gen. 23 N 288.3903** 12.6932** 0.6090** 26.8907** 123.8382** 55.2521 2.0657** 2.9596* 9.6026 6.3770* 1.3884** 0.0024** 154.9414* 0.0007**
S 128.7199** 7.7947** 0.5445 14.1055** 58.2956** 79.1518 2.1971** 5.8633 28.3446 4.4978** 0.9068** 0.0015 132.6449** 0.0009*
Crosses 15 N 267.9802** 13.2764** 0.5472** 26.7468** 117.4804** 71.8387 2.1415** 3.7967* 6.3181 6.5541* 1.4786** 0.0027** 163.1500** 0.0010**
S 146.7558** 8.6764** 0.5336 18.3524** 71.5215** 86.3288 2.7651** 7.6570** 34.3474 4.1724** 1.0627** 0.0017 92.4000 0.0002
Lines 3 N 425.9992** 22.9097** 0.2827 34.6966** 144.5060** 15.5849 0.4753 0.9201 3.4468 10.8234** 2.6980** 0.0047** 363.6389** 0.0010**
S 187.1249** 12.5764** 0.0791 14.7054** 94.6226** 41.0907 0.1743 3.9291 45.6695 7.2201** 0.9444** 0.0003 44.2778 0.0017**
Testers 3 N 593.2675** 29.5764** 1.1601** 81.4050** 328.7918** 75.0582 6.0246** 6.7034** 5.3657 13.0154** 3.73** 0.0065** 115.1389* 0.0017**
S 232.8922** 21.2431 0.9033 55.2759** 207.4118** 102.9634 4.5322** 6.5149 18.5233 9.4407** 2.7032** 0.0038 146.9444 0.0008**
L × T 9 N 106.8781** 4.6319* 0.4311** 5.8775* 38.0347** 89.5169 1.4026* 3.7867* 7.5927 2.9773* 0.3195 0.0008 112.3241 0.0008**
S 104.5873** 3.1875* 0.5619 7.2602** 18.5244** 95.8632 3.0397** 9.8203** 35.8481 1.4003 0.5553** 0.0015 90.2593 0.0002
Parents 7 N 367.1695** 12.0417** 0.8238** 29.8585** 139.5156** 13.1975 1.3576* 1.5231 16.762* 5.9078** 1.3281** 0.0017 149.5655 0.0001
S 108.2244** 6.6607** 0.4883 6.8845** 36.8783** 31.5340 0.9287** 1.6043 8.0127 4.7290** 0.6801** 0.0013 117.6905 0.0001
Crosses vs par. 1 N 443.0883** 8.5069** 0.0312 8.2752** 109.4639** 100.83* 5.8847** 0.4579 8.7517 7.0048s** 0.4568 0.0012 69.4444 0.0003
S 1.6491 2.5069 1.1008 0.9506 9.8282 304.82* 2.5547** 8.7715 80.6254 7.7609** 0.1560 0.0008 841.0000* 0.0051**
Error 46 N 2.2890 1.3146 0.1220 1.5805 15.28 31.8388 0.3762 0.8983 3.7747 0.7248 0.1580 0.0004 39.8545 0.0002
S 2.0609 1.4161 0.2717 1.5567 5.91 49.2103 0.1992 2.7972 32.035 0.8678 0.1909 0.0009 56.0743 0.0002
Whereas Plant height (PH), number of bolls plant
−1
(NB), Boll weight (BW), lint weight (LW), Seed cotton yield plant
−1
(SCY), Lint percentage (L %), Seed index (SI), Lint index (LI), No. of seeds boll
−1
(S/B), Seed mass boll
−1
(SM/B), Lint mass boll
−1
(LM/B)
,
Lint mass per seed, Seed volume per100 seeds (SV) and seed surface area (SSA).
6M. M. ZAFAR ET AL.
Table 4. Mean square values of line × tester analysis for different traits under normal and saline conditions.
SOV DF Trt FF FS FL GOT K
+
/Na
+
Na
+
K
+
H
2
O
2
SOD POD CAT TSP
Rep. 2 N 34.57** 376.96** 1365.40** 877.86** 9.17* 426.43** 1302.73** 0.03** 65.17** 251.94** 948.88** 0.27**
S 44.21** 330.61** 1207.03** 848.72** 0.25** 4120.27** 16081.6** 0.41** 210.22** 358.77** 1767.35** 0.45**
Gen. 23 N 0.40** 10.84** 7.25** 36.07** 3.13** 89.86 396.51** 0.01** 3.62** 42.63** 126.58** 0.06**
S 0.53** 7.92** 9.76** 25.78** 0.59** 138.16** 726.37** 0.01** 30.54* 27.62** 85.68** 0.05**
Crosses 15 N 0.43** 15.49** 9.91** 47.52** 2.63* 31.51 468.59** 0.00 2.73** 43.16** 163.22** 0.06**
S 0.75** 10.17** 13.31** 26.85** 0.54** 101.34** 763.97** 0.01* 13.71 32.57** 99.97** 0.05**
Lines 3 N 1.03** 15.83** 13.72** 116.02** 5.37** 78.83 335.01** 0.00 1.47 83.11** 77.42** 0.06**
S 2.01** 10.08** 24.38** 38.13** 0.15** 134.80* 375.18** 0.02* 4.6709 65.83** 50.26 0.05
Testers 3 N 0.12** 4.80** 11.48** 49.37** 1.83 10.27 890.04** 0.00 0.67 57.05** 108.65* 0.12**
S 0.42** 4.47** 13.1** 11.71 1.32** 193.92** 1564.54** 0.02* 4.4869 40.22** 57.78 0.06
L × T 9 N 0.34** 18.93** 8.12** 24.07** 1.98 22.83 372.63** 0.00 3.84** 25.21** 210.01** 0.04**
S 0.44** 12.11** 9.71** 28.14** 0.41** 59.33 626.72** 0.01 19.8073 18.93** 130.61** 0.05**
Parents 7 N 0.39** 1.64* 1.64 15.11* 4.23** 170.09** 280.93** 0.01** 4.95** 46.95** 65.27** 0.06**
S 0.09** 3.71** 1.87 19.65** 0.77** 231.38** 723.53** 0.00 26.3117 20.95** 64.91 0.05
Crosses vs par. 1 N 0.00 5.55** 6.66** 9.56 2.94 403.37** 124.48** 0.06** 7.58** 4.4065 6.14 0.06*
S 0.31** 3.63 11.78** 52.63** 0.00 37.92 182.16** 0.13** 312.64** 0.05 16.74 0.04
Error 46 N 0.02 0.84 0.90 7.15 1.43 68.18 25.05 0.00 1.45 6.52 28.81 0.01
S 0.03 1.40 1.27 8.47 0.05 48.59 28.67 0.00 16.44 5.77 45.41 45.41
Whereas, FF for Fiber Fineness, FS- Fiber Strength, FL- Fiber Length, GOT- Ginning Out Turn, K
+
/Na
+
Ratio, Na
+-
Sodium, K
+
- Potassium, H
2
O
2
- Hydrogen Per Oxide, SOD- Superoxide dismutase, POD-
Peroxidase CAT- Catalase, TSP, Total Soluble Protein
JOURNAL OF NATURAL FIBERS 7
Table 5. General combining ability (GCA) effects of all parental genotypes in normal and salt stress (15 ds m-1).
Lines Trt PH NB BW LW SCY L% SI LI S/B SM/B LM/B LM/S SV SSA
FH-114 N −7.12** −1.98** −0.13 −2.48** −5.05** −1.52 −0.11 −0.40 0.00 −1.36** −0.68 −0.03** −1.42 −0.01*
S −5.44** −1.48** −0.12 −3.95** 2.64 0.00 0.83 0.00 −2.60 −1.12** −0.40** −1.42 −1.42 −1.50**
NS-131 N −0.50ns 0.19 0.06 −0.27ns 0.58 1.1 0.06 0.23 −0.28
ns
0.12 0.07 0.00 −1.67 0.00
S −0.50ns 0.44 0.05 −1.20ns 1.14 0.17 −0.09 0.01 −0.35 0.38 0.07 −1.67 −1.67 0.27
MS-71 N 7.45** 0.60 0.19 1.20** 2.63** −0.16 0.25 1.13 0.75 0.41 0.19 0.00 0.67 0.00
S 4.01** 0.85* 0.05 2.56** −1.42 −0.10 −0.34 0.00 1.21 0.62* 0.25 0.67 0.67 1.16**
IUB-65 N 0.16 1.19** −0.12 1.01 1.84* 0.58 −0.20 1.04 −0.46 0.82** 0.42** −0.20** 2.42 1.01**
S 0.92 * 0.19 0.02 0.20 −0.08 −0.07 −0.40 0.00 1.74 0.11 0.09 2.42 2.42 0.08
S.E N 0.4666 0.3542 0.0986 0.4113 0.7525 1.8277 0.1795 0.2944 0.5978 0.2593 0.1270 0.0065 2.3686 0.0044
S 0.4117 0.3847 0.1636 0.7020 2.3671 0.1397 0.5733 0.0042 1.9058 0.2843 0.1376 2.3686 2.3686 0.3251
CRS-2007 N 2.88** 1.69** 0.31** 2.99** 5.32** 2.57 0.49* 0.95** −0.44 1.05** 0.66** 0.03** −4.92* 0.00
S 1.91** 1.52** 0.36* 4.64** 3.68 0.37* 1.01 0.01** −0.87 0.93** 0.62** 0.02* −4.92* 2.78**
CIM-573 N 9.87** −1.65** −0.32** −2.40** −5.56** 1.11 −1.01** −0.55** 0.88 −1.16** −0.50** −0.02** 3.17 −0.01
S −6.24** −1.65** −0.27 −5.09** 0.81 −0.88** −0.61 −0.02** 1.75 −1.15** −0.52** −0.02** 3.17 −2.29**
FH-312 N 6.50** −0.98** −0.21** −1.94** −3.27** −3.28 −0.03 −0.64* −0.60 −0.57* −0.42** −0.01* 0.17 −0.00
S 3.93** −0.31 −0.14 −1.24 −1.38 0.04 0.08 0.00 −0.90 −0.15 −0.16 0.00 0.17 −0.83*
Kahkasahn N 0.49 0.94* 0.22* 1.36** 3.50** −0.39 0.49* 0.24 1.15 0.68* −0.26* 0.01 1.58 0.01**
S 0.41 0.44 0.06 1.69* −3.10 0.47** −0.48 0.00 0.01 0.38 −0.06 −0.00 1.58 0.34
SE N 0.4666 0.3542 0.0986 0.4113 0.7525 1.8277 0.1795 0.2944 0.5978 0.2593 0.1270 0.0065 2.3686 0.0044
S 0.4117 0.3847 0.1636 0.7020 2.3671 0.1397 0.5733 0.0042 1.9058 0.2843 0.1376 0.0098 2.3686 0.3251
Plant height (PH), number of bolls plant
−1
(NB), Boll weight (BW), lint weight (LW), Seed cotton yield plant
−1
(SCY), Lint percentage (L %), Seed index (SI), Lint index (LI), No. of seeds boll
−1
(S/B), Seed
mass boll
−1
(SM/B), Lint mass boll
−1
(LM/B)
,
Lint mass per seed, Seed volume per100 seeds (SV) and seed surface area (SSA).
8M. M. ZAFAR ET AL.
Table 6. General combining ability (GCA) effects of all parental genotypes in normal and salt stress (15 ds m-
1
).
PARENTS FF FS FL GOT K
+
/Na
+
Na
+
K
+
H
2
O
2
SOD POD CAT TSP
LINES Trt
FH-114 N −0.03 −0.58 * −1.04 ** 1.96 ** −0.34 1.76 2.82 * −0.02 0.45 3.11 ** −3.28 0.02
S −0.04 −0.95 * −1.70 ** −1.10 0.14 −2.01 4.44 ** −0.02 0.10 2.64 ** 0.08 0.04
NS-131 N 0.21 ** −0.40 1.15 ** −3.69 ** 0.78 * −2.98 4.78 ** 0.01 0.07 0.72 2.32 0.08 *
S −0.03 0.29 1.29 ** −1.49 −0.11 4.98 * 1.93 0.00 −0.70 0.90 0.28 0.01
MS-71 N 0.22 ** 1.71 ** 0.66 * 3.11 ** −0.73 2.48 −7.27 ** 0.03 −0.12 −0.64 1.66 −0.00
S 0.53 ** 1.14 ** 1.06 ** 2.47 * −0.08 −1.97 −8.19 ** −0.04 0.79 −0.66 2.31 0.05
IUB-65 N −0.41 ** −0.73 ** −0.77 ** −1.38 0.28 −1.26 −0.34 −0.01 −0.39 −3.19 ** −0.70 −0.09 *
S −0.46 ** −0.48 −0.66 * 0.13 0.05 −0.99 1.82 0.06 * −0.20 −2.88 ** −2.67 −0.10 *
S.E N 0.0427 0.2402 0.2640 0.7097 0.3758 2.5901 1.2852 0.0258 0.3835 0.6204 1.6665 0.0373
S 0.0502 0.3526 0.3186 0.9794 0.0676 2.1169 1.6019 0.0249 1.2457 0.7587 2.3390 0.0413
TESTERS
CRS-2007 N 0.15 ** 0.92 ** 1.00 ** 1.63 * −0.39 1.05 −9.75 ** 0.01 −0.06 −1.34 * 1.12 −0.15 **
S 0.20 ** 0.71 1.16 ** 1.23 −0.28 ** 2.74 −10.57 ** 0.03 0.18 −1.06 2.11 −0.10 *
CIM-573 N −0.07 −0.29 0.07 −0.96 −0.26 0.50 −3.02 * 0.01 −0.31 1.52 * −3.32 0.04
S −0.25 ** 0.04 0.30 −1.13 −0.23 ** 3.72 −5.97 ** −0.07 * −0.88 1.98 * −2.89 0.00
FH-312 N −0.01 −0.12 −1.34 ** −2.40 ** 0.45 −0.65 10.68 ** −0.02 0.20 −2.34 ** −1.40 0.05
S −0.01 0.02 −1.35 ** −0.31 0.44 ** −5.00 * 15.59 ** 0.02 0.16 −1.97 * −0.42 0.02
Kahkashan N −0.07 −0.52 * 0.27 1.73 * 0.20 −0.89 2.09 −0.01 0.17 2.16 ** 3.59 * 0.06
S 0.07 −0.78 * −0.12 0.21 0.07 −1.46 0.95 0.01 0.54 1.06 1.20 0.08
S.E N 0.0427 0.2402 0.2640 0.7097 0.3758 2.5901 1.2852 0.0258 0.3835 0.6204 1.6665 0.0373
S 0.0502 0.3526 0.3186 0.9794 0.0676 2.1169 1.6019 0.0249 1.2457 0.7587 2.3390 0.0413
Whereas, FF for Fiber Fineness, FS- Fiber Strength, FL- Fiber Length, GOT- Ginning Out Turn, K
+
/Na
+
Ratio, Na
+-
Sodium, K
+
- Potassium, H
2
O
2
- Hydrogen Per Oxide, SOD- Superoxide dismutase, POD-
Peroxidase CAT- Catalase, TSP, Total Soluble Protein
JOURNAL OF NATURAL FIBERS 9
under saline condition, NS-131 indicated positive and significant results whereas FH-312 showed
negative significant estimates. For K
+
conc. line FH-114 and tester FH-312 indicated positive and
significant GCA effects under normal and salt stress conditions. Most of the lines and tester exhibited
nonsignificant GCA effects for H
2
O
2
and SOD. Line FH-114 and tester CIM-573 found good general
combiner in both conditions with significant and positive estimates for POD. All lines and testers
presented nonsignificant results under both conditions for CAT except Kahkashan, which showed
positive and significant estimates under normal (3.59) conditions.
Specic combining ability eects under normal and saline conditions
Crosses FH-114× CIM-573, FH-114× FH-312, MS-71× FH-312, and NS-131× CRS-2007 showed
positive and significant SCA effects for plant height under normal and saline conditions (Tables 7
and 8). For number of bolls per plant, only cross FH-114× Kahkashan showed positive and significant
SCA effects in both conditions. All crosses showed non-significant SCA effects for individual boll
weight except FH-114× FH-312 under saline conditions. FH-114× Kahkashan showed positive and
significant SCA effects for lint weight in both conditions. FH-114× CIM-573 and FH-114× Kahkashan
showed significant and positive SCA effects under both conditions for seed cotton yield. Only cross
FH-114× Kahkashan showed positive SCA effects for seed index in normal condition whereas under
the saline condition all crosses showed non-significant SCA effects for seed index except FH-114× FH-
312. For lint index under normal condition, FH-114× CIM-573, MS-71× CIM-573, and FH-
114× Kahkashan showed positive SCA effects whereas only FH-114× FH-312 in saline condition
showed significant and positive SCA effects. For seed mass per boll, FH-114× FH-312 displayed
positive and significant estimates in saline conditions. FH-114× Kahkashan, NS-131× CIM-573 and
FH-114× FH-312 showed positive and significant specific combining ability for seed volume in normal
and saline conditions, respectively. Cross FH-114× CRS-2007 showed significant positive SCA effects
for seed surface area under normal conditions whereas IUB-65× CIM-573 and FH-114× CIM-573
exhibited positive SCA effects and IUB-65× kahkashan showed negative SCA effects in saline condi-
tions. MS-71 × CRS-2007 indicated positive and significant SCA effects for fiber fineness, fiber
strength, fiber length under both conditions. MS-71 × CRS-2007 indicated positive and significant
SCA effects for GOT in normal conditions. Only MS-71× FH-312 presented significant and positive
SCA under the salt condition for K
+
/Na
+
ratio. MS-71× Kahkashan showed positive and significant
SCA effects for catalase and total soluble proteins under normal conditions. For Na
+
Only MS-
71× Kahkashan exhibited positive and significant SCA in saline conditions. IUB-65× Kahkashan,
MS-71× CRS-2007, and MS-71× FH-312 had good SCA in both conditions for K
+
. For SOD NS-
131× CIM-573 exhibited non-significant SCA under normal conditions. IUB-65× CRS-2007 were
found to be a good specific combiner for POD in both conditions. Cross MS-71× Kahkashan were
identified as a good specific combiner for both CAT and TSP in normal condition whereas under salt
condition only for TSP.
Estimation of gene action by variance of GCA and SCA of dierent traits of various genotypes
of cotton under normal and saline conditions
All the traits of various genotypes of cotton showed a variance of SCA higher than the variance of GCA
in normal condition except H
2
O
2
while in saline condition all traits showed a variance of SCA higher
than the variance of GCA except thenumber of seeds per boll, lint mass per plant, and seed surface area
(Tables 9 and 10). When the Variance of SCA is higher than the variance of GCA, it confirms the
predominance of dominant gene action hence depicts that heterosis breeding can be successful for this
program.
10 M. M. ZAFAR ET AL.
Table 7. Specific combining ability (SCA) effects of all crosses in normal and salt stress.
Crosses Trt PH NB BW LW SCY L% SI LI S/B SM/B LM/B LM/S SV SSA
FH-114× CRS2007 N −5.28 ** 0.06 −0.28 −0.67 −1.74 2.11 −1.05 ** −0.55 1.15 0.02 0.07 0.00 4.71 −0.02 **
S −4.03 ** −0.10 −0.38 −1.53 * −2.04 −1.80 2.89 −1.80 0.08 0.08 −0.16 −0.02 −2.83 −0.01
FH-114× CIM-573 N 6.95 ** 0.73 0.36 1.86 * 3.38 * 2.89 0.91 * 1.25 * −1.81 0.40 0.32 0.02 −7.54 0.03 **
S 5.70 ** 0.73 0.06 0.90 2.98 * −0.74 −1.75 −0.74 0.60 0.60 0.12 0.00 −4.25 0.02 *
FH-114× FH-312 N 1.73 1.36* 0.08 0.73 4.05 * −9.39 * 0.73 * −1.16 1.74 1.03 0.06 0.00 4.46 0.00
S −0.72 1.73* 0.79* 2.38 ** 5.86* 3.42 ** 1.12* 3.42 ** 1.24 1.24* 0.69* 0.04 9.75 * −0.01
FH-114× KAHKASHAN N −3.41 ** 1.85 * 0.57* 1.91 * 5.69 ** 7.39 0.75* 1.46 * 1.08 1.45 ** 0.85* −0.01 6.63 0.00
S −0.95 −1.35 −0.27 −1.74 * −3.80 * −0.88 −0.02 −0.88 −0.93 −0.93 −0.45 −0.02 −2.67 0.00
MS-71× CRS-2007 N 1.21 −1.44 0.18 −1.25 −2.55 −1.07 0.28 0.07 0.69 −0.99 −0.50 −0.03 1.04 0.00
S −0.10 −1.02 0.36 −0.66 −0.40 0.15 1.13 0.15 −0.65 −0.65 −0.42 −0.01 −0.25 0.01
MS-71× CIM-573 N −0.14 −0.77 0.45 * 0.63 −0.19 5.61 0.00 1.22 * 1.07 −0.74 −0.10 0.00 1.13 0.00
S −0.93 −0.85 0.46 0.31 −0.36 1.96 −2.68 1.96 −0.73 −0.73 −0.19 0.00 −0.33 0.01
MS-71× FH-312 N 2.96 ** 0.23 −0.52 * −0.68 −1.92 0.34 −0.55 −0.46 −2.21 0.14 0.13 0.01 1.13 −0.01
S 4.51 ** 0.48 −0.64 −0.84 −1.52 −1.98 1.31 −1.98 n 0.43 0.43 0.11 0.00 −3.67 −0.01
MS-71× KAHKASHAN N −4.03 ** 1.98 ** −0.11 1.29 4.66 ** −4.87 0.27 −0.84 0.45 1.59 ** 0.46 0.02 −3.29 0.01
S −3.47 ** 1.40 −0.18 1.18 2.28 −0.13 0.24 −0.13 0.96 0.96 0.49 0.02 4.25 −0.01
NS-131× CRS-2007 N 4.82 ** 0.48 0.15 1.32 2.35 0.60 0.44 0.63 −1.08 0.30 0.18 0.01 −4.79 −0.01
S 3.19 ** 0.56 0.15 1.52 * 1.78 0.92 −0.81 0.92 0.25 0.25 0.32 0.01 3.08 0.00
NS-131× CIM-573 N −6.29 ** 0.15 −0.53 * −1.35 −2.13 −2.85 −0.56 −1.12 −0.70 0.21 −0.02 0.00 10.29 * −0.02 *
S −3.36 ** −0.27 −0.42 −1.40 * −1.89 −0.57 −2.74 −0.57 0.00 0.00 −0.25 −0.01 −3.00 0.00
NS-131× FH-312 N −7.33 ** −0.85 0.27 −0.15 −1.43 3.85 0.10 0.85 0.33 −0.74 −0.16 −0.01 −7.71 0.01
S −8.84 ** −0.94 0.08 −1.10 −1.43 −0.93 2.16 −0.93 −0.61 −0.61 −0.37 −0.02 −1.67 0.00
NS-131× KAHKASHAN N 8.79 ** 0.23 0.11 0.18 1.21 −1.60 0.01 −0.36 1.46 0.24 0.00 −0.01 2.21 −0.01
S 9.02 ** 0.65 0.19 0.99 1.54 0.58 1.39 0.58 0.36 0.36 0.30 0.01 1.58 −0.01
IUB-65× CRS-2007 N −0.75 0.90 −0.06 0.60 1.94 −1.64 0.33 −0.15 −0.76 0.67 0.25 0.02 −0.96 0.01
S −0.95 0.56 −0.13 0.67 0.67 0.72 −3.21 0.72 0.33 0.33 0.25 0.02 0.00 0.00
IUB-65× CIM-573 N −0.52 −0.10 −0.28 −1.14 −1.05 −5.65 −0.36 −1.36 * 1.44 0.14 −0.20 −0.02 −3.88 0.00
S −1.40 0.40 −0.10 0.20 −0.72 −0.64 7.17 −0.64 0.13 0.13 0.31 0.01 7.58 −0.03 **
IUB-65× FH-312 N 2.63 ** −0.44 0.17 0.10 −0.70 5.21 −0.29 0.77 0.14 −0.43 −0.03 0.00 2.12 −0.01
S 5.05 ** −0.27 −0.03 −0.43 0.08 −0.51 −2.34 −0.51 −0.07 −0.07 −0.23 −0.01 −4.42 0.01
IUB-65× KAHKASHAN N −1.36 −0.35 0.17 0.43 −0.19 2.08 0.32 0.74 −0.82 −0.37 −0.01 0.00 2.71 0.00
S −4.59 ** −0.69 0.26 −0.43 −0.03 0.43 −1.62 0.43 −0.39 −0.39 −0.34 −0.01 −3.17 0.02 *
S.E N 1.3197 1.0019 0.2778 1.1633 2.1285 5.1695 0.5076 0.8326 1.6909 0.7336 0.3595 0.0183 5.9775 0.0124
S 1.1644 1.0882 0.4627 0.9196 1.9855 6.6953 0.3950 1.6215 5.3905 0.8042 0.3891 0.0278 6.6993 0.0118
Plant height (PH), number of bolls plant
−1
(NB), Boll weight (BW), lint weight (LW), Seed cotton yield plant
−1
(SCY), Lint percentage (L %), Seed index (SI), Lint index (LI), No. of seeds boll
−1
(S/B), Seed
mass boll
−1
(SM/B), Lint mass boll
−1
(LM/B)
,
Lint mass per seed, Seed volume per100 seeds (SV) and seed surface area (SSA).
JOURNAL OF NATURAL FIBERS 11
Table 8. Specific combining ability (SCA) effects of all crosses in normal and salt stress.
CROSS Trt FF FS FL GOT KNA NA K HO SOD POD CAT TSP
MS-71× Kahkashan N −0.02 0.00 −0.38 −1.12 −0.71 2.64 −10.30 ** 0.01 0.69 1.09 11.97 ** 0.16 *
S −0.04 −0.15 −0.22 1.31 −0.59 ** 8.78 * −17.58 ** −0.03 2.18 0.13 7.59 0.25 **
FH-114× Kahkashan N −0.15 6.72 * −0.67 −1.25 −0.17 0.27 −3.37 0.01 −0.89 0.70 −8.72 * −0.04
S −0.24 * 0.15 −0.88 1.89 0.11 −4.93 −0.84 −0.02 1.94 1.78 −9.16 −0.06
IUB-65× Kahkashan N −0.10 0.43 1.28 * 0.57 0.70 −2.20 13.58 ** 0.02 0.51 −1.18 3.77 −0.11
S −0.02 −0.04 −0.88 1.37 0.37 * −1.99 14.36 ** 0.01 −2.10 −0.47 3.39 −0.12
NS-131× Kahkashan N 0.27 ** 0.40 −0.23 1.80 0.18 −0.71 0.09 −0.04 −0.31 −0.61 −7.03 * −0.00
S 0.30 ** 0.04 −0.79 −4.57 * 0.12 −1.87 4.05 0.04 −2.02 −1.45 −1.82 −0.08
MS-71× CRS-2007 N 0.28 ** 2.91 ** 1.67 ** 2.99 * 0.20 0.33 8.81 ** −0.01 0.89 −4.23 ** −6.17 0.10
S 0.49 ** 2.18 ** 1.34 * −1.66 0.35 * −3.24 14.45 ** −0.04 3.13 −3.58 * −5.17 0.04
FH-114× CRS-2007 N 0.24 ** 1.69 ** 0.74 2.17 −0.18 0.59 −1.84 −0.00 0.22 −2.53 * −2.63 0.01
S 0.31 ** 1.20 −0.17 −1.26 −0.21 3.95 −5.73 −0.01 −1.87 −2.11 −0.01 0.02
IUB-65× CRS-2007 N 0.22 * −0.66 −0.07 −1.14 0.14 −0.57 1.33 −0.04 0.56 3.46 ** 8.21 * 0.01
S −0.06 −0.12 0.18 2.30 0.05 −0.34 2.45 0.07 −0.79 3.18 * 5.89 −0.03
NS −131 × CRS-2007 N −0.74 ** −3.95 ** −2.34 ** −4.02 ** −0.15 −0.35 −8.30 ** 0.05 −1.68 * 3.30 * 0.59 −0.12
S −0.74 ** −3.26 ** −1.35 * 0.62 −0.20 −0.36 −11.17 ** −0.02 −0.47 2.50 −0.71 −0.03
MS-71× FH-312 N −0.23 * −3.36 ** −2.29 ** 1.25 1.40 −5.55 12.88 ** −0.06 −0.52 −0.26 −3.49 −0.11
S −0.27 * −2.68 ** −2.65 ** 2.43 0.47 ** −4.40 17.72 ** 0.01 −2.55 1.08 −3.71 −0.14
FH-114 × FH-312 N 0.04 1.45 ** 1.03 −3.13 * −0.02 0.78 −0.81 −0.01 0.88 0.68 9.03 * 0.05
S 0.14 0.78 2.15 ** −2.75 0.03 −0.84 1.10 0.04 0.00 −0.62 9.47 0.06
IUB-65× FH-312 N −0.07 −0.76 −0.59 1.74 −1.29 3.51 −17.09 ** 0.05 −0.60 −0.65 −10.09 ** 0.08
S −0.09 −0.41 −1.19 −0.21 −0.46 ** 3.66 −19.12 ** 0.03 0.06 −1.56 −3.80 0.10
NS −131× FH-312 N 0.26 ** 2.68 ** 1.85 ** 0.15 −0.09 1.26 5.02 0.03 0.23 0.22 4.55 −0.02
S 0.21 * 2.32 ** 1.70 * 0.53 −0.04 1.58 0.30 −0.08 2.48 1.10 −1.96 −0.02
MS-71× CIM-573 N −0.03 0.45 1.00 −3.12 * −0.89 2.57 −11.39 ** 0.07 −1.06 3.39 * −2.31 −0.15
S −0.17 0.65 1.53 * −2.09 −0.23 −1.14 −14.59 ** 0.06 −2.77 2.36 1.30 −0.15
FH-114× CIM-573 N −0.14 −2.30 ** −1.11 * 2.22 0.37 −1.64 6.01 * 0.01 −0.22 1.15 2.31 −0.01
S −0.21 * −2.13 ** −1.10 2.12 0.07 1.82 5.47 −0.02 −0.08 0.95 −0.30 −0.02
IUB-65× CIM-573 N −0.04 0.99 * −0.62 −1.17 0.46 −0.74 2.18 −0.04 −0.48 −1.63 −1.89 0.02
S 0.17 0.57 −0.87 −3.46 0.04 −1.34 2.31 −0.11 * 2.83 −1.15 −5.48 0.05
NS −131× CIM-573 N 0.22 * 0.87 0.73 2.07 0.06 −0.20 3.20 −0.04 1.76 * −2.92 * 1.89 0.14
S 0.22 * 0.90 0.44 3.42 0.12 0.65 6.81 * 0.07 0.01 −2.16 4.49 0.12
S.E N 0.1208 0.6795 0.7468 2.0073 1.0629 7.3258 3.6350 0.0729 1.0847 1.7547 4.7135 0.1056
S 0.1419 0.9972 0.9010 2.7701 0.1912 5.9876 4.5309 0.0703 3.5233 2.1458 6.6158 0.1169
Whereas, FF for Fiber Fineness, FS- Fiber Strength, FL- Fiber Length, GOT- Ginning Out Turn, K
+
/Na
+
Ratio, Na
+-
Sodium, K
+
- Potassium, H
2
O
2
- Hydrogen Per Oxide, SOD- Superoxide dismutase, POD-
Peroxidase CAT- Catalase, TSP, Total Soluble Protein
12 M. M. ZAFAR ET AL.
Proportional contribution of lines, testers, and their interactions to the total variances under
normal and saline conditions
The proportional contribution was estimated of lines, testers, and their interaction in the phenotypic
expression of different characters (Tables 9 and 10). Under normal condition, the contribution of
testers found higher than lines and their interaction to the total variance for plant height, while under
saline condition, the contribution of lines and testers interaction found higher than testers and lines.
The contribution of testers was found higher for the number of bolls per plant under normal and
saline conditions. For lint percentage contribution of interaction was found higher in both normal and
saline environments. For seed index, under saline condition, the contribution of L × T was found
higher. Contribution of line × testers interaction was higher for lint index and a number of seeds per
boll for both environments. Testers contributed higher for seed mass per boll and lint mass per boll in
both environments. Under normal condition, the contribution of testers was higher for lint mass
per seed, whilst lines× testers contributed more under saline environment. Contribution of lines found
higher in normal conditions whereas in saline conditions lines × testers interaction contributed higher
for seed volume. The contribution of L × T was found higher for seed density in both environments.
Table 9. Estimation of genetic components and percent contribution of lines, testers, and line × tester for different plant traits under
normal and saline conditions in cotton.
Characters Trt σ2GCA σ2A σ2SCA σ2D
σ2SCA/
σ2GCA
Contribution of
lines
Contribution of
testers
Contribution of
L × T
Plant Height N 5.59 11.18 34.75 34.75 6.21 31.79 25.45 42.76
S 1.46 2.924 34.18 34.18 23.34 25.5 31.70 42.8
No. of bolls/
plant
N 0.30 0.60 1.04 1.04 3.47 34.51 44.55 20.94
S 0.19 0.381 0.47 0.47 2.46 28.99 48.97 22.04
Boll weight N 0.004 0.008 0.1048 0.10 26.2 22.99 59.26 17.75
S −0.001 −0.002 0.08 0.08 −80.3 2.96 33.86 63.18
Lint weight N 0.72 1.44 1.28 1.28 1.76 25.95 60.87 13.18
S 0.38 0.770 1.99 1.99 5.18 16.03 60.24 23.73
Seed cotton
yield
N 2.75 5.51 10.41 10.41 3.77 24.6 55.97 19.43
S 1.84 3.68 4.20 4.20 2.28 26.46 58 15.54
Lint
percentage
N −0.61 −1.22 16.47 16.47 −26.84 8 21 71
S −0.33 −0.66 9.54 9.54 −28.81 9.52 23.85 66.63
Seed index N 0.025 0.05 0.33 0.33 13.17 4.44 56.26 39.3
S −0.009 −0.01 0.93 0.93 −98.44 1.26 32.78 65.96
Lint index N 0.0003 0.0006 0.91 0.91 3.05 4.85 35.31 59.84
S −0.056 −0.112 1.77 1.77 −31.5 10.26 17.02 72.72
No. of seeds/
boll
N −0.04 −0.08 1.10 1.10 −24.86 10.91 16.99 72.1
S −0.052 −0.104 −2.57 −2.57 49.51 26.59 10.79 62.62
Seed mass/
boll
N 0.12 0.24 0.72 0.72 5.82 33.03 39.72 27.25
S 0.09 0.192 0.14 0.14 1.48 34.61 45.25 20.14
Lint mass/boll N 0.04 0.08 0.04 0.04 1.04 36.49 50.54 12.97
S 0.017 0.03 0.10 0.10 6.21 17.77 50.88 31.35
Lint mass/
seed
N 0.0001 0.0002 0.0001 0.0001 1 34.73 47.76 17.51
S 0.000 0.0001 0 0 0 6.26 44.74 49
Seed volume N 1.76 3.52 19.57 19.57 11.09 44.58 14.11 41.31
S 0.074 0.14 7.64 7.64 102.90 9.58 31.81 58.61
Seed surface
area
N 0 0 0.0002 0.0002 0 19.51 33.4 47.09
S 0.000 0 0.0002 0.0002 0 4.73 39.13 56.14
Plant height (PH), number of bolls plant
−1
(NB), Boll weight (BW), lint weight (LW), Seed cotton yield plant
−1
(SCY), Lint percentage (L
%), Seed index (SI), Lint index (LI), No. of seeds boll
−1
(S/B), Seed mass boll
−1
(SM/B), Lint mass boll
−1
(LM/B)
,
Lint mass per seed,
Seed volume per100 seeds (SV) and seed surface area (SSA).
JOURNAL OF NATURAL FIBERS 13
For fiber strength, fiber length, K
+
, K
+
/Na
+
, SOD, CAT, TSP, and hydrogen peroxide, L × T had the
highest contribution under normal and salt stress.
Heterosis
Cross FH-114× Kahkashan exhibited highly significant negative heterotic combinations for plant
height in normal and saline conditions (Tables 11 and 12). For the number of bolls FH-
114× Kahkashan, NS-131× CRS-2007, and MS-71× FH-312 showed positive significant value. Cross
FH-114× Kahkashan and FH-114× FH-312 exhibited the best performance for boll weight and lint
weight under normal and saline conditions, respectively. For seed cotton yield, the hybrid IUB-
65× CIM-573 has a positive significant better parent heterosis value under normal conditions and
MS-71× CRS-2007 has a high significant positive value under saline conditions. For lint percentage,
FH-114× Kahkashan showed positive significant value under normal and saline condition. For seed
index under saline condition FH-114× FH-312, IUB-65× FH-312, and MS-71× CRS-2007 had
significant positive values. For lint index, FH-114× Kahkashan showed a significant and positive
heterotic combination. For seed number per boll, FH-114× Kahkashan and IUB-65× CIM-573 per-
formed better under both normal and saline conditions whereas FH-114× FH-312 depicted positive
heterosis only in saline conditions. For seed volume per seed crosses NS-131× CIM-573 and IUB-
65× CIM-573 were observed as heterotic parent values. FH-114× Kahkashan showed positive and
significant heterosis for fiber fineness under normal conditions whilst IUB-65× Kahkshan and FH-
114× CRS-2007 exhibited positive values under salt conditions. FH-114× Kahkashan and FH-
114× CRS-2007 exhibited positive and significant heterosis for fiber strength under saline and control
conditions, respectively. The highest positive and significant heterosis was observed by MS-71× CRS-
2007 for GOT under control conditions. IUB-65× CIM-573 showed positive and significant heterosis
under both conditions for K
+
/Na
+
whereas MS-71× CRS-2007 and IUB-65× CRS-2007 had positive
and significant heterosis under saline conditions. IUB-65× CIM-573 exhibited negative and significant
Table 10. Estimation of genetic components and percent contribution of lines, testers, and line × tester for different plant traits
under normal and saline conditions in cotton.
Characters Trt σ2GCA σ
2
A σ
2
SCA σ
2
D
σ
2
SCA/
σ
2
GCA
Contribution of
lines
Contribution of
testers
Contribution of
L × T
FF N 0.0032 0.0065 0.1074 0.1074 33.5625 47.22 5.55 47.23
S 0.0108 0.0215 0.1376 0.1376 92.5925 53.41 11.28 35.31
FS N −0.1197 −0.2393 6.0818 6.0818 50.8086 20.44 6.21 73.35
S −0.0672 −0.1343 3.5406 3.5406 −52.6875 19.81 8.78 71.41
FL N 0.0623 0.1245 2.4283 2.4283 38.9775 27.68 23.17 49.15
S 0.1256 0.2512 2.8280 2.8280 30.1715 36.62 19.67 43.71
GOT N 0.8143 1.6286 6.0093 6.0093 7.3797 48.83 20.78 30.39
S −0.0448 −0.0895 5.5447 5.5447 −123.7656 28.4 8.72 62.88
KNA N 0.0225 0.0450 0.0967 0.0967 4.3377 40.81 13.95 45.24
S 0.0046 0.0091 0.1192 0.1192 25.9130 5.7 48.78 45.52
NA N 0.3017 0.6033 −19.2237 −19.2237 −63.7179 50.02 6.52 43.46
S 1.4587 2.9175 1.8534 1.8534 1.2713 26.6 38.27 35.13
K N 3.3319 6.6637 117.6043 117.6043 35.2964 14.3 37.99 47.71
S 4.7659 9.5317 198.6424 198.6424 41.6799 9.82 40.96 49.22
HO N 0.0000 −0.0001 −0.0003 −0.0003 0.00 17.19 11.56 71.25
S 0.0001 0.0003 0.0021 0.0021 21 24.47 28.38 47.15
SOD N −0.0385 −0.0770 0.6939 0.6939 −18.0233 10.78 4.93 84.29
S −0.2115 −0.4230 0.3956 0.3956 −1.8704 6.81 6.54 86.65
POD N 0.6232 1.2464 6.8663 6.8663 11.0178 38.51 26.44 35.05
S 0.4736 0.9471 4.0091 4.0091 8.4651 40.42 24.7 34.88
CAT N −1.6247 −3.2493 58.8957 58.8957 −36.2502 9.49 13.31 77.2
S −1.0637 −2.1274 21.6512 21.6512 −20.3546 10.06 11.56 78.38
TSP N 0.0007 0.0014 0.0086 0.0086 12.2857 19.34 40.05 40.61
S 0.0001 0.0001 0.0115 0.0115 115 18.5 23.23 58.27
Whereas, FF for Fiber Fineness, FS- Fiber Strength, FL- Fiber Length, GOT- Ginning Out Turn, K
+
/Na
+
Ratio, Na
+-
Sodium, K
+
-
Potassium, H
2
O
2
- Hydrogen Per Oxide, SOD- Superoxide dismutase, POD- Peroxidase CAT- Catalase, TSP, Total Soluble Protein
14 M. M. ZAFAR ET AL.
Table 11. Heterosis effects of various plant traits under normal and saline conditions in cotton.
Crosses Trt PH NB BW LW SCY L% SI LI S/B SM/B LM/B LM/S SV SSA
FH-114× CRS2007 N −24.14 * −24.14 * −9.06 −23.51 * −7.12 * 11.83 −21.89 ** −4.14 −2.33 −26.71 * −17.77 −22.22 −13.51 −43.04 **
S −2.61 −18.18 −0.55 −15.41 10.77 * 9.14 −15.30 ** 1.99 3.98 −20.79 −11.92 −19.23 −10.11 −1.21
FH-114× CIM-573 N −33.33 * −33.33 * −8.76 −30.33 * −30.02 ** 15.51 −14.43 * 20.51 −8.27 −36.92 ** −23.38 −18.18 −30.18 ** −37.30 **
S 0.62 −21.43 −3.32 −24.46 −30.98 ** −3.18 −3.54 −8.59 −3.74 −20.86 −22.57 −22.22 0.53 18.77
FH-114× FH-312 N −19.05 −5.56 −13.92 −39.81 ** −21.46 ** −28.51 ** −2.78 −35.80 * −0.77 −13.15 −32.14 * −46.43 * −13.06 −39.30 **
S −8.62 ** 11.76* 29.22* 24.27 51.79 ** 26.52 * 15.88 82.25 ** 8.93 41.53 ** 6.28 33.33 18.09 −27.06 **
FH-114× KAHKASHAN N −53.33 ** 19.05* 38.76 * 48.42 ** 24.45 ** 37.47* 18.31 ** 39.54* 48.27* 55.61 ** 47.61 ** −52.63 ** −30.18 ** −33.26 **
S −11.32 ** −54.17 ** −4.74 50.88 ** 51.76 ** −3.18 −18.28 ** −8.59 2.95 −56.09 ** −49.13 ** −53.33 * −6.44 1.58
MS-71× CRS-2007 N −17.24 −53.33 ** 6.52 −2.04 18.26 ** 9.94 −2.16 17.19 −3.16 −19.98 −11.81 −16.67 −3.30 −11.26 **
S 0.35 −4.55 6.04 16.82 42.86 ** 1.49 7.23 10.56 −2.60 −6.24 −2.91 −7.69 −7.85 8.58
MS-71× CIM-573 N −33.33 ** −17.24 −3.87 −26.62 * −27.85 ** 21.84 * −22.57 ** 14.70 3.07 −37.42 ** −23.90 −25.00 4.06 26.04 **
S −15.31 ** −33.33 * −8.48 −39.18 ** −52.02 ** 10.30 −12.78 * 13.74 −6.70 −33.68 −31.90 −28.57 4.71 27.64 *
MS-71× FH-312 N −12.50 13.33 ** −28.42 ** −36.31 ** −20.70 ** −0.39 −15.87 * −16.33 −14.46 * −11.74 −11.21 3.57 5.08 −39.15 **
S 10.63 ** 11.11 −34.63 ** −35.00 * −9.07 * −12.11 −26.38 ** −41.57 −2.05 −11.16 −1.65 −12.50 −5.24 −27.04 *
MS-71× KAHKASHAN N 6.67 −12.50 −4.53 14.28 −21.89 ** −5.47 −5.09 −7.55 −1.91 6.15 7.33 −5.26 −20.91 ** −29.15 **
S −8.14 ** 4.17 −16.68 8.68 −39.07 * −5.75 −17.18 ** −19.65 −2.58 0.50 12.50 −10.00 3.47 −25.10 *
NS-131× CRS-2007 N 6.90 6.67 −4.21 3.07 −3.88 7.89 0.43 15.76 −10.62 −0.51 7.33 13.89 −11.32 −19.06 **
S 2.52 17.39 22.49 49.79 ** −26.72 ** 20.21 −0.41 41.45 3.34 −5.72 16.52 16.67 7.22 −19.02 *
NS-131× CIM-573 N −31.03 ** 6.90 −36.78 ** −58.82 ** −24.48 ** −3.00 −31.90 ** −35.56 ** −4.71 −32.68 ** −34.88 ** −32.35 * 10.95 −6.06 **
S −20.91 ** −34.78 ** −13.06 −53.22 ** −41.75 ** −3.19 −19.98 ** −15.01 5.91 −41.75 ** −46.61 ** −46.67 10.56 −15.75
NS-131× FH-312 N −34.48 ** −31.03 ** −14.21 * −46.04 ** −19.50 ** 2.10 −9.91 −5.83 −6.26 −37.75 ** −36.43 ** −32.35 * −13.81 −7.09 **
S −13.42 ** −26.09 * 8.67 −30.52 * −28.48 ** −3.65 −1.13 −7.43 14.33 −35.62 ** −37.61 ** −43.33 7.78 18.51 *
NS-131× KAHKASHAN N −6.67 −34.48 ** −7.69 −18.16 * −2.40 −3.57 −5.90 −11.08 0.30 −8.39 −10.53 −21.05 −13.18 −38.90 **
S 9.42 ** 0.00 16.57 11.59 −27.48 ** 5.24 −16.74 ** 10.47 14.87 −12.42 −4.81 −6.67 2.97 −25.66 **
IUB-65× CRS-2007 N 6.25 −6.67 −2.62 6.82 −2.40 2.60 −4.85 7.41 −5.11 4.90 9.51 23.08 0.47 71.45 **
S −3.44 4.17 12.02 38.44 * −0.36 17.98 0.10 36.14 −2.63 −3.90 19.21 12.90 10.53 4.62
IUB-65× CIM-573 N −31.38 ** 6.25 −27.28 ** −53.98 ** 22.17 ** −9.31 −30.24 ** −37.59 ** 8.39 −31.49 ** −37.67 ** −43.59 ** −3.74 −36.46 **
S −20.62 ** −37.50 ** 0.48 −28.06 −38.48 ** 24.29 −45.20 ** −11.94 46.38 * −42.49 ** −24.66 −41.94 38.01 ** 4.62
IUB-65× FH-312 N −31.25 ** −34.38 ** −11.36 −39.55 ** 22.17 ** 4.76 −14.80 * −3.59 −2.27 −31.27 ** −30.19 ** −25.64 5.61 −0.22
S 6.60 ** −29.17 * 21.49 −13.63 3.11 −3.87 10.97 7.42 3.18 −28.93 * −31.88 * −38.71 11.70 −12.02
IUB-65× KAHKASHAN N −12.50 −12.50 1.26 −8.52 −24.68 ** 4.25 −7.74 10.28 8.39 −13.83 −9.71 −5.13 −5.45 25.60 **
S −14.95 ** −25.00 * 17.93 −7.45 −48.80 ** −4.05 −2.51 6.20 0.30 −26.54 * −27.66 −38.71 −1.49 −14.02
Plant height (PH), number of bolls plant
−1
(NB), Boll weight (BW), lint weight (LW), Seed cotton yield plant
−1
(SCY), Lint percentage (L %), Seed index (SI), Lint index (LI), No. of seeds boll
−1
(S/B), Seed
mass boll
−1
(SM/B), Lint mass boll
−1
(LM/B)
,
Lint mass per seed, Seed volume per100 seeds (SV) and seed surface area (SSA).
JOURNAL OF NATURAL FIBERS 15
Table 12. Heterosis effects of various plant traits under normal and saline conditions in cotton.
CROSS Trt FF FS FL GOT K
+
/Na
+
Na
+
K
+
H
2
O
2
SOD POD CAT TSP
MS-71× Kahkashan N 2.00 0.75 −2.41 6.27 −41.01 ** −14.04 −17.28 ** −34.25 −23.75 * −22.07 * 30.90 ** −18.18
S 15.00 ** −5.18 −2.77 6.04 −41.14 ** −1.93 −28.49 ** −38.40 −1.60 −16.89 16.83 16.86
FH-114× Kahkashan N 10.92 ** 4.89 0.91 6.08 −41.75 ** −10.14 −20.56 ** −5.08 −44.21 ** 6.38 −16.65 −79.80 **
S 5.88 * 12.33 ** −0.57 9.47 −28.66 ** −13.63 −25.10 ** −30.40 −5.33 1.47 −22.41 −68.60 **
IUB-65× Kahkashan N −5.36 * 2.67 1.33 4.81 −16.34 −18.08 4.47 −47.87 * −25.37 * −18.30 −9.34 −59.09 **
S 10.19 ** −4.02 0.35 5.33 6.42 −32.35 ** 1.68 −30.30 −46.27 ** −25.44 −5.50 −56.98 **
NS-131× Kahkashan N 0.31 5.05 −4.91 0.65 −34.63 * −20.97 −17.50 ** −16.33 −40.44 ** 8.28 −22.63 * −44.44 *
S 15.66 ** 6.01 −3.31 −16.53 ** −27.37 ** 11.14 −19.54 ** −43.08 * −51.34 ** −4.53 −16.63 −53.49 **
MS-71× CRS-2007 N 1.58 3.22 −1.24 10.92 * 3.83 −20.15 10.61 ** −60.27 * 13.71 −29.95 ** −10.17 44.76
S 13.99 ** −4.16 −6.31 −11.85 * 34.18 ** −18.61 * 15.17 ** −31.93 1.54 −22.10 −15.62 63.41
FH-114× CRS-2007 N 5.07 9.63 ** −0.08 8.85 −15.30 −10.99 −5.23 −35.59 −0.00 −22.55 −7.80 −45.05
S 5.46 * 6.60 −4.32 −8.73 −10.09 −1.22 −10.64 ** −20.17 −19.85 −26.33 * 0.34 −12.79
IUB-65× CRS-2007 N −3.99 −10.68 ** −9.75 ** −7.64 7.59 −16.14 12.15 ** −80.85 ** −13.98 9.30 −10.23 5.98
S −1.10 −13.11 ** −14.87 ** −4.30 37.52 ** −30.44 ** 21.41 ** −10.61 −43.31 ** 2.15 −4.89 −15.65
NS-131 × CRS-2007 N −23.12 ** −21.23 ** −17.37 ** −14.38 ** −12.20 −21.68 −16.42 ** 10.20 −32.41 * 32.96 * −17.15 −41.73
S −1.10 −18.87 ** −15.45 ** −13.10 * −31.11 ** 0.30 −21.20 ** −52.31 ** −47.24 ** 23.38 −18.48 −21.85
MS-71× FH-312 N −16.92 ** −22.52 ** −13.59 ** 1.42 12.39 −38.08 ** 1.78 −79.45 ** −21.41 −41.65 ** −8.03 −47.62
S −13.64 ** −23.12 ** −16.94 ** 3.27 −1.16 −18.79 * −0.82 8.26 −40.76 * −25.38 −17.76 −51.22
FH-114 × FH-312 N −7.48 ** 3.81 1.36 −8.54 −19.54 −17.02 −15.14 ** −48.61 −3.97 −17.07 12.10 −64.86 *
S −4.88 1.76 5.16 −6.34 −28.82 ** −6.51 −18.74 ** 17.54 −27.85 −33.93 19.85 −45.35
IUB-65× FH-312 N −16.51 ** −11.68 ** −12.09 ** −0.20 −26.54 −13.85 −12.39 ** −50.00 * −37.73 ** −33.37 −38.42 ** −3.42
S −11.80 ** −12.41 ** −16.07 ** −0.84 −20.60 ** −24.43 ** −14.97 ** −2.27 −40.61 * −45.48 * −33.33 * −17.39
NS-131× FH-312 N −11.56 ** 3.91 −2.77 −12.59 * −18.75 −24.40 −9.90 ** −31.94 −17.34 −5.86 −5.77 −43.88
S −10.68 ** 5.63 −0.90 −10.69 * −29.28 ** 19.20 −16.32 ** −48.46 * −34.56 * −1.82 −28.53 * −52.94
MS-71× CIM-573 N 1.33 −7.94 ** 3.84 −18.82 ** 37.62 −25.18 −3.44 −17.81 −12.63 −3.62 15.41 −33.33
S 1.89 −6.16 5.13 −18.64 ** −1.65 −6.03 −8.26 * −18.06 −44.64 * 8.47 8.16 −52.38 *
FH-114× CIM-573 N 12.31 ** −13.11 ** 1.07 −7.51 44.50 −28.65 * 0.56 −28.99 −0.63 4.73 18.27 −49.31 *
S −3.41 −11.68 ** 4.26 −7.81 −8.52 5.62 −4.36 −31.25 −30.82 5.51 6.26 −61.22 **
IUB-65× CIM-573 N −4.42 −4.25 −3.01 −23.52 ** 86.44 ** −30.29 * 12.48 ** −72.34 ** −30.94 * −24.84 −18.31 * 0.00
S 3.42 −5.04 −7.90 * −22.53 ** 40.14 ** −23.28 ** 27.94 ** −52.08 ** −29.71 −19.55 −30.44 * −22.45
NS-131× CIM-573 N −0.93 −5.38 −0.15 −13.42 ** 21.39 −30.57 * −7.79 ** −47.83 23.20 −4.68 −3.06 23.61
S 3.47 −1.55 1.55 −10.00 * −28.42 ** 11.12 −11.25 ** −34.03 * −48.74 ** 2.33 −4.95 −10.88
S.E N 0.1281 0.7503 0.7776 2.1838 0.9777 6.7421 4.0872 0.0606 0.9853 2.0857 4.3830 0.1116
S 0.1450 0.9689 0.9230 2.3764 0.1899 5.6919 4.3721 0.0805 3.3114 1.9627 5.5023 0.1073
Whereas, FF for Fiber Fineness, FS- Fiber Strength, FL- Fiber Length, GOT- Ginning Out Turn, K
+
/Na
+
Ratio, Na
+-
Sodium, K
+
- Potassium, H
2
O
2
- Hydrogen Per Oxide, SOD- Superoxide dismutase, POD-
Peroxidase CAT- Catalase, TSP, Total Soluble Protein
16 M. M. ZAFAR ET AL.
heterosis for Na
+
. MS-71× CRS-2007, IUB-65× CRS-2007, and IUB-65× CIM-573 showed positive and
significant heterosis for K
+
concentration in both conditions. IUB-65× CIM-573 presented negative
and significant heterosis for H
2
O
2
under both conditions. Crosses NS-131× Kahkashan and IUB-
65× FH-312 had negative and significant heterosis for SOD. Only NS-131 × CRS-2007 exhibited
positive heterosis for POD under normal conditions. For CAT, only MS-71× Kahkashan presented
positive and significant heterosis under normal conditions.
Discussion
Salt tolerance ability in the cotton plant is a complex quantitative trait and it is the result of various
morphological and yield traits (Shakeel 2017). To push forward breeding program against salinity
tolerance, the effective knowledge about variability, heritability, proportional contribution to the
total variance, gene action, and heterosis is a prerequisite (Tang et al. 1996). Four lines and four
testers were crossed in Line × Tester fashion (4 × 4) and developed 16 F1 hybrids (Ali et al. 2014). It
is pertinent to mention that it provides reliable information about the genetic architecture of
inheritance pattern from its combining ability estimates (Karademir et al. 2009). The proportional
contribution of lines and testers to total variance was variable for all the traits under salt stress
(Ahuja and Dhayal 2007). The results showed that the maximum proportional contribution for
most of the traits under salt stress was contributed by the testers. The argument can be generated on
the earlier findings of many experiments that salt tolerance in the testers is due to the high uptake of
K
+
ions inside the plant that had maintained salt tolerance in the cotton plant (Karademir et al.
2009). Whereas the highest uptake of K
+
ions and balancing ratio of K
+
/Na
+
will lead to the high
salt tolerance ability of the cotton that in turn will increase the seed cotton yield under salt-affected
soils. Production of hydrogen peroxide is considered as a marker in the plant to prepare itself
against the onset of salt stress (Li et al. 2011). SOD is the first line of defense inside the plant cell
against the onset of a high level of ROS species under salt stress. SOD produced in high quantity in
plants with high endurance against salt stress, which indicated that SOD can be considered a valid
parameter to judge the ability of the genotype against salt tolerance (Farooq 2019). Peroxidase
converts hydrogen peroxide into water and oxygen and assists the plant in detoxification mechan-
ism against the ROS species (Farooq et al. 2018). The increase of salt stress directly relates to the
disruption of soluble proteins concentration in the plant cells. It leads to the accumulation of
sodium ions inside the cell membrane that disrupts the internal functions of organelles (Shelke et al.
2017).
General combining ability is due to the additive gene actions whereas the specific combining
ability is due to the non-additive gene actions. FH-114 had highly significant GCA effects for plant
height, bolls per plant, lint weight, seed mass per boll, seed cotton yield, ginning out turn
percentage, and peroxidase. The results suggested that the lines with high GCA effects for specific
traits can be utilized in the breeding program for the development of salt-tolerant genotypes
through hybridization followed by selection breeding (Saleem et al. 2009). Specific combining ability
results indicated that hybrid FH-114 × Kahkashan performed better for agronomic traits under
normal conditions and MS-71× CRS-2007 showed significant SCA effects for fiber traits. FH-114
× FH-312 indicated good specific combining ability under salt stress conditions. Moreover, there
were hybrids in our study with negative values under control and both levels of salt stress which
indicated the existence of different genes with minor effects in each line or preponderance of
epistasis. Non-additive gene action was shown by most of morphological, fiber, ionic, and biochem-
ical traits. High values of nonadditive gene action indicated that the interaction of lines × testers had
the high specific combining ability and fewer genes were responsible for the expression of the
characters (Aslam et al. 2015).
Heterosis study showed that maternal effects had a more pronounced effect on the expression of
morphological, ionic, fiber, and physiological traits. Heterosis was found significantly positive for FH-
114 × KAHKASHAN at the control and 15 dSm
−1
for plant height, a number of bolls, seed cotton
JOURNAL OF NATURAL FIBERS 17
yield, lint weight, lint percentage, lint index, seeds per boll, seed mass per boll, lint mass per boll, and
fiber fineness. FH-114 × FH-312 showed significant heterosis under salt stress for plant height,
a number of bolls, boll weight, seed cotton yield, lint percentage, lint index, seed mass per boll, and
seed surface area. The presence of variation in the performance of the parents and the hybrid
development programs can be attributed to difference in the genetic constitution of the plants and
their specific interaction with the prevailing environment. The presence of nonadditive gene action for
most of the studied traits further suggests that the plant material can be a good option for the hybrid
development. The neighboring countries of Pakistan, India, and China have attained higher cotton
production than Pakistan by developing cotton hybrids (Blaise, Venugopalan, and Raju 2014). The
hybrid development of cotton in Pakistan is at the early stage in Pakistan which needs to be revitalized
to enhance the cotton production in the country.
Conclusions
Salinity severely impacts the capacity of cotton production. The ability of a cotton plant to show
resilience against salt stress is controlled by nonadditive action. Based on information from biome-
trical approaches used herein, FH-114 × KAHKASHAN and FH-114 × FH-312 were found the best
genotypes for future hybrid development. These results are limited to the plant material studied and
therefore, may not be generalized most of the cotton-growing areas facing salinity problem in
Pakistan. Therefore, it is suggested that this information must be substantiated by another genetic
experiment that may involve a reasonable sample of cotton cultivars, evaluated under diverse
environments in order to enhance stress adaptations of our existing commercial cultivars of cotton
and to develop plant material with improved salt tolerance.
Disclosure statement
The authors declare that they have no conflict of interest for the publication of the manuscript.
Funding
Department of Plant Breeding and Genetics, University of Agriculture Faisalabad, Faisalabad-Pakistan and Institute of
Cotton Research, Chinese Academy of Agricultural Sciences, Anyang-China.
Authors’ contributions
MMZ, AR, MAF, and AS conducted experiment, contributed equally, and wrote the initial draft of the manuscript. AR,
HF, and HM contributed in data analysis, MR, MA, and YY proofread the manuscript.
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