Journal of Pure and Applied Agriculture (2023) 8(3): 10-21

ISSN (Print) 2617-8672, ISSN (Online) 2617-8680

http://jpaa.aiou.edu.pk/

Line × tester analysis for yield and its attributed traits in upland cotton
(Gossypium hirsutum L.)
Muhammad Ahmad1*, Asif Saeed1, Khunsa Khakwani2, Muhammad Aqib3, Afaaq Tariq1, Nabeel Yaqoob1, Rujab
Nadeem1, Ayesha Bashir4 and Muhammad Saad Rafique4
1
Department of Plant Breeding and Genetics, University of Agriculture, Faisalabad, 38000, Faisalabad, Pakistan
2
Cotton Research Station, Ayub Agriculture Research Institute, Faisalabad, 38000, Faisalabad, Pakistan
3
Institute of Botany, University of Punjab, Lahore, 54000, Punjab, Pakistan
4
Department of Plant Breeding and Genetics, The Islamia University, Bahawalpur, 63100, Bahawalpur, Pakistan

*Corresponding author: Muhammad Ahmad (ahmaduaf2013@gmail.com)

Abstract

Cotton production in our country is declining due to produced cultivars that are not well-adapted to changing environmental
conditions. New high-yielding, climate-resilient cotton genotypes are needed to enhance our economy. The objective of the
study was to assess the effects of gene action, combining ability and heterosis on seed cotton yield and its associated traits.
Five lines and three testers of upland cotton (Gossypium hirsutum L.) were used to produce fifteen F1 hybrids using a line ×
tester mating design. The research was conducted in a randomized complete block design with three replications at Cotton
Research Station, Ayub Agricultural Research Institute (AARI), Faisalabad in the Kharif season of 2020-21. The mean squares
of all the traits were found significant. The results showed that among the female lines; FH-414 performed best for plant
height, days to 50% flowering, boll weight, fiber strength, and fiber length while FH-490 was found best for seed cotton yield
and GOT%. Testers concluded that EYE-111 was an excellent general combiner in terms of plant height, monopodial
branches, days to maturity, bolls per plant, days to 50% flowering, seed cotton yield, and fiber quality. The cross combinations
i.e., FH-492 × CIM-602 for plant height, FH-414 × CIM-602 for fiber fineness and fiber length, and FH-492 × NIAB-SANAB-
M for seed cotton yield showed good SCA effects. FH-414 × CIM-602 and FH-415 × EYE-111 depicted maximum heterosis
for fiber traits and seed cotton yield, respectively. FH-ANMOL × EYE-111 had shown maximum heterosis for sympodial
branches and days to 50% flowering, FH-414 × CIM-602 for GOT% and fiber fineness. all the characters were regulated by
non-additive kind of gene action. The above-mentioned genotypes having GCA and SCA effects could be further exploited for
hybrid/variety development programs to cope with unexpected climatic conditions.

Keywords: Combining ability, Cotton revival, GCA, Heterosis, Heterobeltoisis, SCA

To cite this article: Ahmad, M., Saeed, A., Khakwani, K., Aqib, M., Tariq, A., Yaqoob, N., Nadeem, R., Bashir, A., &
Rafique, M. S. (2023). Line × tester analysis for yield and its attributed traits in upland cotton (Gossypium hirsutum L.).
Journal of Pure and Applied Agriculture, 8(3), 10-21.

Introduction was due to the use of poor-quality seeds, inadequate water

availability, fluctuation in weather during flowering (Singh et
Cotton is the world‟s most important fiber crop and is al., 2022) and less water available during the crucial plant
commonly known as silver fiber (Ali et al., 2012; Zia et al., development stages (Government of Pakistan [GOP], 2021-
2015; Munir et al., 2018). The upland cotton (G. hirsutum 22). It is very important to develop high-yielding and well-
L.) is generally known as “American cotton” and is adapted cultivars under climate change scenarios to fulfill
cultivated in about 90% of the area throughout the globe future requirements (Shuli et al., 2018; Shoukat et al., 2020;
(Wang et al., 2020). Cotton is widely grown in China, Shaukat et al., 2021; Arif et al., 2022).
India, the United States, the Middle East and Australia Cotton breeders are constantly using different breeding
(Tantuway & Patil, 2020). Cotton is said to be the methods to develop high fiber-yielding varieties by utilizing
backbone of Pakistan‟s economy (Zia et al., 2018a; Zia et diversified genetic resources of cotton (Khokhar et al., 2018).
al., 2022; Abbas et al., 2022; Khan et al., 2023). It However, it is important to understand the various cotton
contributed 0.8% to the GDP of the country and 4.1% of components that increase yield before beginning any breeding
the total value added to agriculture (Government of effort to boost seed cotton yield (Nizamani et al., 2017,
Pakistan [GOP], 2021-22). Since Pakistan's independence, Soomro, 2020). A thorough understanding of yield components
scientists have studied and developed new varieties with like bolls per plant, boll weight, lint percentage etc. and how
the best traits, yet per-unit seed cotton yield is still poor they are inherited is critical for making informed breeding
(Zia et al., 2018b; Makhdoom et al., 2019). The overall decisions. The knowledge of combining ability, obtained
area of cotton has increased, but the production remained through biometrical tools like line x tester analysis, provides
below as compared to last year. The decrease in production key insights into the gene action governing yield and
Muhammad Ahmad et al Journal of Pure and Applied Agriculture (2023) 8(3): 10-21
component traits. Information on general combining ability 490, FH-492 and FH-ANMOL, and three testers, NIAB-
(GCA) helps identify parents with superior breeding value, SANAB-M, CIM-602 and EYE-111, all belonging to upland
while specific combining ability (SCA) indicates hybrid cotton (G. hirsutum L.) were collected from Cotton Research
combinations with optimum trait expression. Such Station, AARI, Faisalabad. During October 2019-20, the
knowledge empowers breeders to choose ideal parents and parental genotypes were grown in earthen pots of size “30×30
crosses for harnessing both additive and non-additive cm2” and placed in the glasshouse. At flowering, these lines
genetic variances. This enables more rapid genetic gains crossed in line×tester fashion. The emasculation was done in
from selection and hybridization. Therefore, a good grasp the evening and the buds were covered with soda straw to
of combining abilities and inheritance patterns is pivotal avoid foreign contamination while the cotton flower was also
before initiating any breeding endeavor to improve seed tagged in the evening, which would be used as a male pollen
cotton yield (Khokhar et al., 2018). parent the next day. On the next day, emasculated buds were
A biometrical tool called line × tester analysis provides pollinated using selected male flowers and covered again.
details on the variances in combining ability and the effects Selfing was also done in some of the buds by using nail polish
of the genotypes (Sajjad et al., 2016) regarding parent (sticky material) to prevent opening. To get sufficient seed, the
genotypes and their crosses which could be useful for a maximum number of crosses were made between the lines and
future breeding program (Abdel-Aty et al., 2022). Plant testers following the emasculation and pollination procedures
researchers widely used line × tester for early generation described by Khan et al. (2018). According to Khan et al.
selection (Usharani et al., 2016; Sivia et al., 2017). GCA (2018), making the maximum number of crosses with proper
aids in the identification of potential parental genotypes for emasculation technique is crucial for obtaining enough F 1
hybridization and subsequent selection in a population that hybrid seed for replicated field trials. The list of the parental
segregates, whereas SCA is necessary for hybrid seed genotypes along with their F1 crosses is shown in Table 1.
production and non-additive gene action is more The F1 crossed seed of 15 hybrids and parental genotypes (5
significant as compared to additive gene action (Manan et lines, 3 testers) were planted in the field under randomized
al., 2022). GCA variation was greater for seed index and complete block design with three replications during the Kharif
GOT% whereas SCA was greater for plant height, boll season of 2020-21. For recording the observations, three plants
weight, bolls per plant and seed cotton yield (Zafar et al., from each replication were randomly tagged. The distance
2022). The study helped to identify the suitable parents and between rows (R×R) and plants (P×P) was maintained as 75
their crosses after evaluating their combining ability (GCA and 45cm, respectively. The recommended agronomic/cultural
and SCA) for yield and its attributing traits. practices e.g., fertilizers, weeding, chemicals and irrigations,
The objectives of the study were to develop single etc. were followed as per recommendation and crop
cross hybrids in upland cotton (Gossypium hirsutum L.) requirement of Punjab Agriculture Department [PAD] (2022),
using line x tester mating design and evaluate them along Government of Punjab throughout the growing season. The
with parents in a field trial. The study aimed to assess the data were recorded for the parameters viz., plant height (cm),
gene action governing yield, yield components, and fiber boll weight (g), days to maturity, days to 50% flowering,
quality traits in the breeding materials. Combining ability number of bolls per plant, monopodia per plant, sympodia per
effects, including general combining ability (GCA) and plant, seed-cotton-yield (kg/ha), ginning outturn percentage
specific combining ability (SCA), were estimated for all (GOT%) and various fiber related traits such as fiber
the traits to understand the relative importance of additive fineness/micronaire value (µg/in), fiber length (mm) and fiber
and non-additive gene actions. The overarching goal was strength (g/tex) were measured using computerized high
to identify superior parents and cross combinations that volume instrument developed by Uster Spinlab Company
could be further exploited to improve yield potential, yield (model number HVI 900) that provides a comprehensive
components and fiber quality in cotton breeding programs profile of raw data. The variance analysis was conducted by
tailored to current climatic conditions. using Steel et al. (1997) approach to study the variability
among the genotypes/crosses. GCA and SCA of the hybrids
Materials and Methods and parents were calculated through line-by-tester analysis
developed by Kempthorne (1957). Further, the heterosis and
The research was performed at the experimental site of the heterobeltiosis were estimated through the formula suggested
Cotton Research Station (CRS), Ayub Agricultural by Falconer and Mackay (1996). All statistical analysis was
Research Institute (AARI), Faisalabad (31°26'0"N performed using R-Studio software (version 4.2.0) with the
73°4'0"E). The five parental lines, FH-414, FH-415, FH- package „Agricolae‟ for line×tester analysis.

Table 1 Parental (male and female) upland cotton genotypes and their F1 hybrids
Lines (Female parent) Crosses (F1 hybrids)
1. FH-414 1. FH-414 × NIAB-SANAB-M
2. FH-415 2. FH-415 × NIAB-SANAB-M
3. FH-490 3. FH-490 × NIAB-SANAB-M
4. FH-492 4. FH-492 × NIAB-SANAB-M
5. FH-ANMOL 5. FH-ANMOL × NIAB-SANAB-M
11
Muhammad Ahmad et al Journal of Pure and Applied Agriculture (2023) 8(3): 10-21

6. FH-414 × CIM-602
Testers (Male parent)
7. FH-415 × CIM-602
8. FH-490 × CIM-602
1. NIAB-SANAB-M 9. FH-492 × CIM-602
2. CIM-602 10. FH-ANMOL × CIM-602
3. EYE-111 11. FH-414 × EYE-111
12. FH-415 × EYE-111
13. FH-490 × EYE-111
14. FH-492 × EYE-111
15. FH-ANMOL × EYE-111

Results and Discussion for plant height, monopodial branches, sympodial branches,
days to 50% flowering, days to maturity, boll weight, bolls per
Variance analysis and estimation of gene action plant, seed cotton yield, fiber strength and fiber length (Rani et
al., 2020). When the GCA/SCA ratio is negative and smaller
Analysis of variance (Line × Tester) depicted that than one, it is indicative of "additive x dominant" gene
genotypic differences for treatments (eight parents and interactions (Falconer & Mackay, 1996). The ratio between the
fifteen crosses) were found significant (P<0.01) for all the GCA and SCA variance estimates for studied characteristics
traits under study (Table 3). Kempthorne‟s approach suggested that "additive dominance" gene interactions were
(1957) of line-tester was used for further data analyses. present for plant height, GOT%, and fiber length. Previous
The results revealed that parents, parents vs. crosses and studies have documented various gene influences on the
interaction (line × tester) were significant while non- characteristics studied. The dominant gene effects on ginning
significant differences were found in lines and testers. The outturn fiber length and strength were reported (Nagarajan et
significant parents and their crosses (F1) indicated that the al., 2022).
traits were governed by non-additive as well as additive Bolls per plant is a reliable predictor of the overall
gene action. The interaction between lines and testers also production of seed cotton. Higher values for the SCA variance
had a favorable effect on the differences in results. The component suggested that the number of bolls produced per
findings revealed significant genetic variations between the plant was the primary trait where non-additive gene activity
lines and the testers. The results indicated that hybrid trait was evident. Heterosis breeding is indicated to improve the
expression would likely to differ based on parental number of flowers developed by each plant. Non-additive gene
combinations (Farooq et al., 2022). The contribution of action was reported in bolls per plant (Fatima et al., 2022),
lines, testers and their crosses (F1 hybrids) are given in monopodia per plant (Kanasagra et al., 2022) and sympodia
Table 2. Estimation of GCA (σ²gca) and SCA (σ²sca) per plant, SCY (Imran et al., 2012) and fiber strength (Balci et
variance and expressing them as σ²gca/σ²sca may be used al., 2023). Characters should not be selected that are influenced
to evaluate the significance of GCA and SCA. If this ratio by non-additive gene action until their genes have been well-
is around 1, it shows the significance of additive gene established in subsequent generations (Aydin et al., 2019;
action (Mawblei et al., 2022). Tables 4 & 5 display GCA Salman et al., 2019). Boll weight (Manan et al., 2022), fiber
and SCA variations for yield and fiber-related length and fiber fineness (Fatima et al, 2022) and GOT%
characteristics, respectively. A higher magnitude of σ²sca (Manan et al., 2022) were governed by additive gene action.
than σ²gca was found, indicating non-additive gene action

Table 2 Contribution (%) of lines, testers and their crosses (line × tester)
PH MB SB DFF DOM NBP BW SCY GOT FF FS FL
Line 6.11 50.96 55.48 79.35 31.94 10.36 48.08 56.56 18.58 62.52 14.68 29.87
Tester 13.65 0.19 0.56 0.85 20.28 4.58 14.35 3.28 6.93 17.69 19.74 2.68
Line × tester 80.24 48.85 43.95 19.80 47.78 85.05 37.57 40.16 74.49 19.79 65.58 67.45
PH: Plant height; MB: Monopodial branches; SB: Sympodial branches; NBP: Number of bolls per plant; DFF: Days to fifty percent
flowering; DOM: Days to maturity; BW: Boll weight; SCY: Seed cotton yield; GOT%: Ginning out turn; FF: Fiber fineness; FS: Fiber
strength; FL: fiber length

Combining ability additive gene activity was evident. Heterosis breeding is

indicated to improve the number of flowers developed by each
Bolls per plant is a reliable predictor of the overall plant. Non-additive gene action was reported in bolls per plant
production of seed cotton. Higher values for the SCA (Fatima et al., 2022), monopodia per plant (Kanasagra et al.,
variance component suggested that the number of bolls 2022) and sympodia per plant, SCY (Imran et al., 2012) and
produced per plant was the primary trait where non- fiber strength (Karademir et al., 2016). Characters should not

12
Muhammad Ahmad et al Journal of Pure and Applied Agriculture (2023) 8(3): 10-21

be selected that are influenced by non-additive gene action General Combining ability of testers
until their genes have been well-established in subsequent
generations (Salman et al., 2019; Aydin et al., 2019). Boll Among the testers, CIM-602 showed positive GCA effects on
weight (Manan et al., 2022), fiber length and fiber fineness plant height (5.96**), monopodial branches (0.02), sympodial
(Fatima et al, 2022) and GOT% (Manan et al., 2022) were branches (0.58**) and GOT% (0.30*) and fiber strength
governed by additive gene action. (0.95**) while negative GCA depicted for seed cotton yield (-
35.67**). EYE-111 had negative significant GCA effects for
Combining ability plant height (-4.04**), monopodial branches (-0.04*), days to
50% flowering (-0.64**) and days to maturity (-1.6**) while
The predominance of non-additive gene action observed in depicting maximum positive GCA results for the number of
this study aligns with findings from several previous bolls per plant (2.60), seed cotton yield (25.33**) and fiber
investigations on cotton (Kumar et al., 2017). Max et al. fineness (0.16**). NIAB-SANAB-M depicted maximum
(2021) favored heterosis breeding over pure line selection positive significant results for days to 50% flowering (0.82**),
due to the preponderance of dominance and epistasis effect days to maturity (1.80**), boll weight (0.01**) and
and highlighted reciprocal recurrent selection as an monopodial branches (0.02) while negative GCA effect for
efficient long-term breeding strategy to accumulate fiber fineness (-0.10*). The same kind of results was reported
favorable epistatic interactions. Kumar et al. (2017) earlier in the research of Kumar et al., (2017) and Thiyagu et
advised rigorous multi-environment testing and delayed al., (2019). EYE-111 as a tester was a good general combiner
selection in advanced generations to deal with for plant height, monopodial branches, days to 50% flowering,
unpredictable performance of segregating populations. days to maturity, number of bolls per plant, seed cotton yield
Taken together, the complex non-additive genetic and fiber fineness. The same kind of results were reported
architecture necessitates tailored breeding schemes, earlier in the research of Ahuja and Dhayal (2007). NIAB-
advanced molecular tools and extensive field-testing to SANAB-M is the best tester for boll weight. CIM-602 could be
enhance selection efficiency and maximize genetic gains. used as a good general combiner for the number of sympodia,
GCA and SCA mean squares showed additive and non- GOT% and fiber strength. Jatoi et al. (2011) reported the same
additive gene action (Table 3). The general combining result. All three testers (NIAB-SANAB-M, CIM-602 and
ability of parents (lines and testers) (Table 4) and SCA EYE-111) showed non-significant GCA effects for fiber
effects (Table 5) of F1 hybrids for yield and fiber-related length.
traits were also calculated.
Specific combining ability
General Combining ability of lines (GCA line)
All the crosses showed the highly positive significance of SCA
Among lines, FH-414 depicted highly significant and effects. Positive but significant SCA effects depicted by FH-
positive GCA effects for days to maturity (3.0**), average 490×CIM-602 for plant height (15.71**), FH-414×NIAB-
boll weight (0.20**), fiber length (0.54**), fiber strength SANAB-M for boll weight (0.24**), FH-490×EYE-111 for
(0.65**) while maximum negative GCA effect showed for days to maturity (4.20**), FH-ANMOL×NIAB-SANAB-M for
plant height (-2.98**), bolls per plant (-7.60**) and days to days to 50% flowering (7.51**), FH-ANMOL×CIM-602 for
50% flowering (-7.11**). FH-415 had the highest bolls per plant (15.27**), FH-492×NIAB-SANAB-M for seed
significant and negative GCA effects for days to maturity cotton yield (203.0**) and fiber strength (2.72**), FH-
(2.00**), monopodial branches (-0.56**), number of ANMOL×EYE-111 for number of sympodial branches
sympodial branches (-6.49**), GOT% (-1.14**) and fiber (6.58**), FH-415×EYE-111 for GOT% (2.49**) and FH-
fineness (-0.31**) while FH-ANMOL showed maximum 414×CIM-602 (1.09**) for fiber length. These results were
positive and highly significant GCA effects number of matched with Zafar et al. (2022). While maximum negative
bolls per plant (3.73**), number of sympodia (6.29**). SCA effect was revealed by FH-492×CIM-602 for plant height
FH-490 showed maximum positive GCA effects for plant (-17.62**), FH-414×EYE-111 for boll weight (-0.23**) and
height (5.36**), number of monopodia (0.89**), SCY fiber fineness (0.21*), FH-492×EYE-111 for days to maturity
(154.07**), GOT% (0.90**), fiber fineness (0.30**) while (-4.80**) and fiber strength (-1.98**), FH-ANMOL×EYE-111
negative GCA recorded for average boll weight (-0.27**), for days to 50% flowering (-4.02**), FH-415×CIM-602 for
fiber strength (-1.10**). Negative GCA is desirable for monopodia per plant (-0.91**). These results were similar to
plant height, days to maturity and days to fifty percent Usharani et al. (2016). As per the requirement for days to
flowering to develop short to medium-heightened, short- maturity and days to 50% flowering, there is a need to find out
duration and early maturing genotypes (Munir et al., 2018). if the cross has negative combining ability effects (Khokhar et
FH-414 from lines is a good general combiner for plant al., 2018). Best Specific combiners are FH-492×CIM-602 for
height, boll weight, days to 50% flowering, fiber fineness plant height, FH-492×EYE-111 for boll weight, FH-
and fiber strength while FH-415 is the best general ANMOL×EYE-111 for days to 50% flowering and number of
combiner for monopodia per plant. FH-490 for GOT% and sympodial branches FH-ANMOL×CIM-602 for bolls per
FH-ANMOL for bolls per plant, sympodia per plant plant, FH-415×CIM-602 for number of monopodial branches,
showed as a best general combiner. FH-492×NIAB-SANAB-M for fiber strength and could be
13
Muhammad Ahmad et al Journal of Pure and Applied Agriculture (2023) 8(3): 10-21

used in further breeding programs. The same kind of FH-414×CIM-602 for GOT% and fiber length could be used in
results were reported earlier in the research of Munir et al. the future for breeding purposes.
(2018).
Estimation of heterobeltiosis (%)

Estimation of heterosis (%) Heterobeltiosis (better parent heterosis) for various yields and
fiber-related character was given in Table 7. Maximum
A broad range of heterosis from positive to negative for positive heterobeltiosis was depicted by the crosses, FH-
most of the characters was found in the crosses of upland 490×CIM-602 for plant height (13.06%), FH-492×EYE-111
cotton as shown in Table 6. Maximum but highly for boll weight (13.28%) and monopodial branches (91.03%),
significant and positive heterosis was found in the crosses FH-414×NIAB-SANAB-M for days to maturity (3.84%), FH-
such as FH-490×CIM-602 for plant height (14.11%), FH- 492×NIAB- SANAB-M for days to 50% flowering (8.14%),
492×EYE-111 for boll weight (13.83%), FH-414×NIAB- FH-ANMOL×CIM-602 bolls per plant (92.37%), sympodia
SANAB-M for days to maturity (4.02%), FH-492×NIAB- per plant (83.33%) and fiber strength (13.28%), FH-415×EYE-
SANAB-M for days to 50 percent flowering (10.84%), FH- 111 for seed cotton yield (32.94%), FH-ANMOL×EYE-111
ANMOL×CIM-602 for bolls/plant (108.26%) and fiber for GOT% (7.01%), FH-490×EYE-111 for fiber fineness
strength (13.98%), FH-ANMOL×EYE-111 for sympodial (1.75%) and FH-414×CIM-602 for fiber length (6.98%). While
branches (104.11%) and seed cotton yield (25.83%), FH- maximum negative but significant heterobeltiosis was shown
414×CIM-602 for GOT% (8.32%) and fiber length by FH-490×EYE-111 for plant height (-11.43%), FH-
(84.33%) and FH-490×EYE-111 for fiber fineness 415×EYE-111 for boll weight (-14.30%), FH-492×EYE-111
(3.69%). Maximum negative and significant heterosis was for sympodia per plant, days to maturity and seed cotton yield
shown by FH-490×EYE-111 for plant height (-10.33%), (-16.67%, -3.07%, -10.26%, respectively), FH-ANMOL×EYE-
FH-414×EYE-111 for boll weight (-5.46%), FH- 111 for days to 50% flowering (-12.42%), FH-492×NIAB-
492×EYE-111 for days to maturity (-2.82%), SANAB-M for GOT%. (-7.49%) and FH-414×CIM-602 for
ANMOL×EYE-111 for days to 50 percent flowering (- fiber fineness (-85.04%). Negative heterobeltiosis is desirable
8.89%), FH-490×NIAB-SANAB-M for bolls/plant (- for plant height (Fetahu et al., 2015), the number of monopodia
20.87%) for bolls per plant, FH-ANMOL×CIM-602 for the (Monicashree et al., 2017; Riaz et al., 2023). The crosses
number of monopodial branches (100.00%), FH- having minimum values for days to 50% flowering would be
415×CIM-602 for the number of monopodial branches (- used as selection criteria for early maturity. Similar results of
33.33%), FH-492×CIM-602 for seed cotton yield (- positive heterobeltiosis were reported for boll weight (Rani et
3.77%), FH-415×NIAB-SANAB-M (-6.77%) for GOT%, al., 2020), sympodia per plant, SCY (Munir et al., 2018),
FH-414×CIM-602 for fiber fineness (-74.29%), FH- GOT% (Islam et al., 2021), fiber length and fiber fineness
492×EYE-111 fiber strength (-8.23%) and FH- (Vadodariya et al., 2022), fiber strength (Hamed and Said,
ANMOL×NIAB-SANAB-M for fiber length (-9.44%). The 2021). The crosses having maximum values for the number of
most important aim of the study is to find hybrids that have bolls per plant was the best hybrid and could be used in the
short to medium-heightened plants that prevent lodging future for breeding purpose. Similar outcomes for positive
losses and as a result yield increases. The crosses having heterobeltiosis for bolls per plant (Keerthivarman et al., 2022).
minimum plant height (Zhang et al., 2017), days to The best desirable hetrobeltoisis showed by hybrids such as
maturity and days to 50% flowering (Khokhar et al., 2018) FH-490×EYE-111 for plant height and fiber fineness, FH-
would be used as selection criteria for early maturity, short 492×EYE-111 for boll weight and days to maturity, FH-
duration, lodging resistant genotypes while positive ANMOL×EYE-111 for days to 50% flowering and GOT%,
heterosis is required for bolls per plant, bolls per plant, and FH-ANMOL×CIM-602 for sympodial branches, bolls per
sympodia per plant (Zapadiya et al., 2021). Monopodial plant and fiber strength, FH-414×CIM-602 for fiber length and
branches bear indirect fruits, so a smaller number of FH-415×EYE-111 for monopodial branches and seed cotton
monopodia was required. Similar findings were also yield.
reported by Khokhar et al. (2018) for negative heterosis of
monopodia per plant while positive heterosis is required Conclusion
for bolls per plant (Zafar et al., 2022), fiber length
(Thiyagu et al., 2019) fiber fineness and fiber strength Estimation of combining ability effects plays a key role in
(Yehia & El-Hashash, 2019). Thiyagu et al. (2019) determining yield and related traits. The SCA effect showed
reported similar results of positive heterosis for seed cotton non-additive gene influences in all characteristics studied. To
yield while Ahuja (2018) reported GOT%. Maximum prevent losing better genotypes, selection might be delayed for
desirable heterosis was found by crosses like FH- future generations. F1 crosses contributed more than lines or
490×EYE-111 for plant height, FH-492×EYE-111 for days testers in terms of plant height, bolls per plant, days to
to maturity and boll weight, FH-ANMOL×EYE-111 for maturity, GOT%, fiber strength and fiber length, but lines
number of sympodial branches, FH-415×CIM-602 for consistently performed better in terms of monopodial and
number of monopodia, FH-490×NIAB-SANAB-M for sympodial branches, days to fifty percent flowering, boll
bolls per plant, FH-415×EYE-111 for seed cotton yield and weight, seed cotton yield and fiber fineness. The tester is
14
Muhammad Ahmad et al Journal of Pure and Applied Agriculture (2023) 8(3): 10-21

positioned between lines and crosses for all attributes. One would be useful before varietal release to stabilize hybrid
of the lines, FH-414 had the highest GCA for plant height, performance. The best hybrids identified here, like FH-492 ×
monopodial branches, days to 50% flowering, boll weight, NIAB-SANAB-M and FH-414 × CIM-602, need further
fiber strength and fiber length whereas FH-490 displayed evaluation across environments and assessment for yield
the highest GCA for fiber fineness, seed cotton yield, stability before commercial exploitation. The best general
GOT% and days to maturity. EYE-11 had the best GCA combiners, namely FH-414, FH-490 and EYE-111, should be
effect for plant height, monopodial branches, days to 50% extensively utilized in cotton breeding tailored to current
blooming, and seed cotton among the tested varieties. The climatic conditions. The insights gained from this study
study has some potential limitations that need to be regarding gene action, general combining ability, specific
considered. non-additive gene action governed most traits, combining ability and heterosis will be valuable for selecting
indicating performance may fluctuate in subsequent ideal parents and crosses to develop high-yielding cotton
generations. Specific combining ability was also varieties adapted to changing climate.
significant, highlighting the influence of specific parental
combinations. Future multi-location and multi-year trials Disclosure of competing interest: No potential conflict of interest
across contrasting environments are recommended to was reported by the author(s).
validate the present findings. Expanding the genetic base
by including more lines and testers could help test a wider Acknowledgments: This research didn‟t receive a specific grant from
any funding agency in the public, commercial or not-for-profit
array of combinations. Advanced generation progeny tests sectors.

15
Muhammad Ahmad et al Journal of Pure and Applied Agriculture (2023) 8(3): 10-21

Table 3 Variance analysis of combining ability for fiber quality and yield contributing traits in upland cotton
SOV DF PH MB SB DFF DOM NBP BW GOT% SCY FF FS FL
81.565* 0.010*
Replications 2 235.406** 0.406 33.783 86.623 115.928* 0.060 0.0138 0.016
* * 1988.484 1.328 *
1.477* 166.979* 113.286* 24.300* 569.825* 0.227*
Treatments 22 400.026** 7.717** 72.615** 7.349**
* * * * * * 7547.998** 81.456 **
12.232* 0.316* 206.343* 243.963*
Parents (P) 7 361.714** 0.262 15.851** 72.857** 47.143** 5.385** 4.419**
* * 609.963** * *
7.548* 1847.33* 16.984* 5222.68* 0.010* 24.917* 73662.143* 149.973*
P vs. C 1 171.602** 5.845** 2.365**
* * * * * * * * 60.544**
1.651* 122.517* 141.175* 30.857* 498.819* 0.199*
Crosses (C) 14 435.498** 7.654** 0.225** 9.167**
* * * * * * 6294.577** 1.695**
392.056*
Lines (L) 4 93.189 2.944 237.922 34.500 180.922 0.334 4.978 0.493* 4.710
* 1103.884 1.772
Testers (T) 2 416.089 0.022 4.822 8.422 43.800 160.067 0.200 3.711 1435.193 0.279 12.665 0.318
1.411* 25.800* 742.456* 0.131* 10.520*
L×T 1 611.506** 94.239** 48.922** 9.978 0.078*
* * * * 10104.77** * 2.001 **
Error 44 0.588 0.224 0.722 0.108 0.202 0.200 0.001 0.243 3.143 0.027 0.226 0.266

σ²gca -6.22 0.01 1.00 3.26 0.18 -8.61 0.00 -0.08 693.09 0.01 -0.05 -0.01
σ²sca 203.74 0.39 31.31 16.28 8.58 247.45 0.04 3.24 15440.64 0.01 3.40 0.61
σ²gca/σ²sca -0.03 0.02 0.03 0.20 0.02 -0.03 0.06 -0.03 0.04 0.36 -0.01 -0.02
* = Significant (P≤0.05), ** = Highly significant (P≤0.01) ns = Non-significant; SOV: Sources of variance; DF: Degree of freedom; PH: Plant height; MB: Monopodial branches; SB:
Sympodial branches; NBP: Number of bolls per plant; DFF: Days to fifty percent flowering; DOM: Days to maturity; BW: Boll weight; SCY: Seed cotton yield; GOT%: Ginning out turn;
FF: Fiber fineness; FS: Fiber strength; FL: fiber length; σ²gca: general combining ability variance; σ²sca: specific combining ability variance 0

Table 4 Predicted general combining abilities (GCA) effects for lines and testers concerning yield, yield components and fiber traits
Genotypes PH MB SB NBP DFF DOM BW SCY GOT% FF FS FL
Lines
FH-414 -2.98** -0.33* -3.82** -7.60** -7.11** 3.00** 0.20** 31.73** 0.18 0.06 0.65** 0.54**
FH-415 0.36 -0.56** -6.49** 0.73** -4.44** -2.00** 0.08** 58.07** -1.14** -0.31** -0.15 0.23
FH-490 5.36 ** 0.89** 1.18** 2.84** 3.56** -0.33** -0.27** 154.07** 0.90** 0.30** -1.10** -0.45**
FH-492 -1.31** -0.22 2.84** 0.29* 9.44** 0.67** 0.12** -112.60** -0.17 0.10 -0.06 -0.47**
FH-ANMOL -1.42** 0.22 6.29** 3.73** -1.44** -1.33** -0.13** -31.27** 0.23 -0.14* 0.65** 0.15
SE 0.26 0.16 0.28 0.15 0.11 0.15 0.00 0.26 0.16 0.05 0.16 0.17
Tester
NIAB-SANAB-M -1.91** 0.02 -0.56** -3.67** 0.82** 1.80** 0.01** 10.33** -0.57** -0.10 * -0.07 -0.05
CIM-602 5.96** 0.02 0.58** 1.07** -0.18* -1.60** 0.11** -35.67** 0.30* -0.06 0.95** 0.16
EYE-111 -4.04** -0.04 -0.02 2.60** -0.64** -0.20** -0.12** 25.33** 0.27* 0.16** -0.88** -0.11
SE 0.20 0.12 0.22 0.12 0.08 0.12 0.00 0.20 0.13 0.04 0.12 0.13
PH: Plant height; MB: Monopodial branches; SB: Sympodial branches; NBP: Number of bolls per plant; DFF: Days to fifty percent flowering; DOM: Days to maturity; BW: Boll weight;
SCY: Seed cotton yield; GOT%: Ginning out turn; FF: Fiber fineness; FS: Fiber strength; FL: fiber length
16
Muhammad Ahmad et al Journal of Pure and Applied Agriculture (2023) 8(3): 10-21

Table 5 Predicted specific combining abilities (SCA) effects of crosses for yield, yield components and fiber traits
Genotypes PH MB SB NBP DFF DOM BW SCY GOT% FF FS FL
FH-414 × NSM 6.91** 0.20 3.89** 10.00** -3.16** -0.13 0.24** -36.33** 0.30 -0.05 -1.59** -1.25**
FH-414 × CIM-602 -15.96** -0.47 -3.24** -8.73** -0.16 -0.73** -0.01** -101.33** 1.11** -0.16 -0.69* 1.09**
FH-414 × EYE-111 9.04** 0.27 -0.64 -1.27** 3.31** 0.87** -0.23** 137.67** -1.41** 0.21* 2.28** 0.16
FH-415 × NSM -6.42** 0.42 5.56** -0.33 -0.82** 0.87** -0.14** -62.67** -1.92** -0.14 -0.29 0.44
FH-415 × CIM-602 5.71** -0.91** -1.58** -6.07** 1.18** -0.73** 0.21** 5.33 -0.56 0.01 -0.19 -0.07
FH-415 × EYE-111 0.71** 0.49 -3.98** 6.40** -0.36* -0.13 -0.06** 57.33** 2.49** 0.13 0.48 -0.37
FH-490 × NSM -1.42** -0.02 -0.11 -24.44** -3.82** -2.80** -0.10** -39.67** 1.22** 0.13 0.96** 0.44
FH-490 × CIM-602 15.71** -0.02 4.76** 10.49** 2.18** -1.40** -0.09** 85.33** -0.11 0.03 0.14 -0.63**
FH-490 × EYE-111 -14.29** 0.04 -4.64** 13.96** 1.64** 4.20** 0.19** -45.67** -1.11** -0.15 -1.10** 0.18
FH-492 × NSM 5.24** 0.09 -0.78* 12.78** 0.29 1.20** -0.17** 203.00** -1.50** -0.07 2.72** 0.88**
FH-492 × CIM-602 -17.62** 0.76* -1.91** -10.96** 0.29 3.60** -0.03** -51.00** 1.26** 0.13 -0.74* -0.63**
FH-492 × EYE-111 12.38** -0.84** 2.69** -1.82** -0.58** -4.80** 0.20** -152.0** 0.24 -0.06 -1.98** -0.25
FHA × NSM -4.31** -0.69* -8.56** 2.00** 7.51** 0.87** 0.17** -64.33** 1.91** 0.13 -1.79** -0.51*
FHA× CIM-602 12.16** 0.64* 1.98** 15.27** -3.49** -0.73** -0.08** 61.67** -1.69** 0.00 1.48** 0.24
FHA× EYE-111 -7.84** 0.04 6.58** -17.27** -4.02** -0.13 -0.10** 2.67 -0.22 -0.13 0.31 0.27
SE 0.44 0.27 0.49 0.26** 0.19 0.26 0.00 0.44 0.28 0.11 0.27 0.30
NSM: NIAB-SANAB-M; FHA: FH-ANMOL; PH: Plant height; MB: Monopodial branches; SB: Sympodial branches; NBP: Number of bolls per plant; DFF: Days to fifty percent flowering;
DOM: Days to maturity; BW: Boll weight; SCY: Seed cotton yield; GOT%: Ginning out turn; FF: Fiber fineness; FS: Fiber strength; FL: fiber length

Table 6 Heterosis for yield and fiber-related traits in upland cotton

Crosses PH MB SB NBPP DFF DOM BW SCY GOT% FF FS FL
FH-414 × NSM 3.79** 23.08 39.74** 58.25** 5.43** 4.02** 10.53** 73.13** 1.36 -73.97** -6.16** 60.22**
FH-414 × CIM-602 -8.05** 0.00 17.42** 14.15** -7.49** 1.39** 1.14** 19.07* 8.32** -74.29** 0.54 84.33**
FH-414 × EYE-111 1.81 ** 33.33 20.50** 45.81** -5.90** 2.42** -5.46** 34.44** 1.13 -70.50** 4.22** 76.32**
FH-415 × NSM -5.77** 23.08 30.86** 40.18** -1.54** 2.19** -4.08** 33.98* -6.78** -17.20** 1.46 -0.01
FH-415 × CIM-602 4.52** -33.33 9.32** 33.91** -1.53** -0.96** 2.37** 33.06* 1.40 -11.84** 5.74** 2.07
FH-415 × EYE-111 -4.43** 33.33 -5.39* 77.38** -5.02** -0.43* -5.39** 48.83** 8.28** -2.49 1.44 -0.18
FH-490 × NSM 0.84* 46.67** 49.33** -20.87** 1.00** 0.00 -1.91** -3.87 6.13** 0.68 2.53* -6.24**
FH-490 × CIM-602 14.11** 57.14** 74.50** 77.97** 4.97** -1.55** -4.48** 72.24** 7.81** 0.66 3.49** -6.32**
FH-490 × EYE-111 -10.33** 57.14** 29.03** 98.24** 2.45** 1.54** 2.59** 88.64** 4.42** 3.69 -7.58** -4.74**
FH-492 × NSM 12.57** 23.08 47.44** 59.18** 10.48** 2.33** 6.62** 68.88** -5.31** -10.02** 11.01** -4.31**
FH-492 × CIM-602 -0.46 66.67** 48.39** 9.96** 8.40** 1.29** 7.54** 30.42 6.21** -3.84 2.69* -5.79**
FH-492 × EYE-111 13.69** -16.67 57.76** 40.50** 5.52** -2.82** 13.83** 67.13** 2.86** -1.13 -8.23** -5.68**
FH-A × NSM -0.88* 16.67 44.68** 63.21** 6.29** 1.65** 6.84** 71.17** 8.32** -3.39 -1.52 -9.44**
FH-A × CIM-602 12.59** 100.00** 95.71** 108.26** -6.56** -1.47** -2.97** 100.79** 4.21** -4.24 13.98** -3.59**
FH-A × EYE-111 -6.48** 63.64** 104.11** 28.23** -8.89** -0.95* -4.56** 42.31** 7.14** 0.04 3.19* -4.60**
NSM: NIAB-SANAB-SANAB; FH-A: FH-ANMOL; PH: Plant height; MB: Monopodial branches; SB: Sympodial branches; NBPP: Bolls per plant; DFF: Days to fifty percent flowering;
DOM: Days of maturity; BW: boll weight; SCY: Seed cotton yield; GOT%; Ginning out turn percentage; FF: Fiber fineness; FS: fiber strength; FL: Fiber length

17
Muhammad Ahmad et al Journal of Pure and Applied Agriculture (2023) 8(3): 10-21

Table 7 Heterobeltiosis for yield and fiber-related traits in upland cotton

Crosses PH MB SB NBPP DFF DOM BW SCY GOT% FF FS FL
FH-414 × NSM 3.46 ** 14.29 31.32** 45.54** -11.40** 3.84** 1.22** 66.75** 0.46 -84.80** -11.26** -7.77**
FH-414 × CIM-602 -9.77 ** 0.00 9.64** 2.54** -11.18** 1.04** -2.44** 15.34* 4.57** -85.04** -5.13** 6.98**
FH-414 × EYE-111 0.21 33.33 16.87** 35.78** -10.87** 1.54** -13.43** 28.85** -1.64 -82.94** -1.54 2.27
FH-415 × NSM -8.92 ** 14.29 19.10** 40.18** -6.51** 2.10** -13.11** 19.25** -7.04** -19.49** 1.35 -3.20*
FH-415 × CIM-602 3.25 ** -33.33 -1.12 30.51** -7.35** -1.56** -2.38** 26.31** -1.52 -13.28** 5.61** 1.81
FH-415 × EYE-111 -5.88** 33.33 -11.27** 75.00** -11.8** -1.54** -14.30** 31.88** 5.96** -3.08 1.44 -0.67
FH-490 × NSM -2.24 ** 37.50* 45.45** -22.88** -1.63** -1.03** -2.05** -5.88 5.90** -0.85 2.42 -7.04**
FH-490 × CIM-602 13.06** 37.50* 68.83** 77.97** 1.28** -2.06** -9.37** 64.15** 4.78** 0.29 3.36* -9.83**
FH-490 × EYE-111 -11.43** 37.50* 28.21** 90.68** -2.48** 1.54** 2.44** 83.78** 2.26* 1.75 -7.59** -8.11**
FH-492 × NSM 4.13** 14.29 38.55** 46.62** 8.14** 1.02** 6.11** 56.34** -7.49** -10.74** 9.66** -4.55**
FH-492 × CIM-602 -9.77** 66.67** 38.55** 3.76** 5.11** 0.51** 2.36** 29.15* 1.12 -5.72 1.22 -8.79**
FH-492 × EYE-111 3.35** -16.67 53.01** 27.82** 0.93** -3.07** 13.28** 53.99** -1.34 -5.16 -9.43** -8.47**
FH-A × NSM -2.17** 0.00 39.73** 54.46** 4.56** 0.86** 3.88** 66.02** 6.42** -9.99** -1.89 -12.10**
FH-A × CIM-602 8.73** 83.33** 90.28** 92.37** -8.95** -1.72** -5.52** 82.34** 3.28** -9.80** 13.28** -9.10**
FH-A × EYE-111 -9.41** 50.00* 91.03** 22.94** -12.42** -1.20** -7.21** 38.73* 7.01** -3.72 2.69 -9.86**
NSM: NIAB-SANAB-M; FH-A: FH-ANMOL; PH: Plant height; MB: Monopodial branches; SB: Sympodial branches; NBPP: Number of bolls per0plant; DFF: days to fifty
percent0flowering; DOM: Days to maturity; BW: boll weight; SCY: Seed cotton yield; GOT%; Ginning out turn percentage; FF: Fiber fineness; FS: fiber strength; FL: fiber length.

18
Muhammad Ahmad et al Journal of Pure and Applied Agriculture (2023) 8(3): 10-21
References and their cross combinations for yield and fiber
associated attributes in American cotton (Gossypium
Abbas, M. A., Sarwar, G., Shah, S. H., Muhammad, S., hirsutum L.). Biological and Clinical Sciences Research
Zafar, A., Manzoor, M. Z., Murtaza, G., & Latif, M. Journal, 2(1), 1-12.
(2022). Evaluating the effectiveness of boron on the Fetahu, S., Aliu, S., Rusinovci, I., Zeka, D., Shabani, H. P.,
growth of cotton (Gossypium hirsutum L.) under Kaul, H. P., & Elezi, F. (2015). Determination of
saline conditions. Pakistan Journal of Botany, 54(5), heterosis and heterobeltiosis for plant height and spike
1619-1628. grain weight of F1 generation in bread wheat.
Abdel-Aty, M. S., Youssef-Soad, A., Yehia, W. M. B., El- International Journal of Ecosystems and Ecology
Nawsany, R. T. E., Kotb, H. M. K., Ahmed, G. A., Science, 5, 431-436.
Hasan, M. E., Salama, E. A., Lamlom, S. F., Saleh, F. Government of Pakistan [GOP]. (2021-22). Economic survey
H., & Shah, A. N. (2022). Genetic analysis of yield of Pakistan 2020-21. Ministry of Economics, Finance
traits in Egyptian cotton crosses (Gossypium Division, Pakistan, 2, 17-41.
barbdense L.) under normal conditions. BMC Plant Hamed, H. H., & Said, S. R. N. (2021). Estimation of heterosis
Biology, 22(1), 462. https://doi.org/10.1186/s12870- and combining ability for yield and fiber quality traits by
022-03839-8 using line x tester analysis in cotton (Gossypium
Ahuja, S. L. (2018). Heterosis studies for high ginning barbadense L.). Menoufia Journal of Plant Production, 6,
outturn percent and related traits in Gossypium 35-51. https://doi.org/10.21608/mjppf.2021.169080
hirsutum cotton. International Journal of Agricultural Imran, M., Shakeel, A., Azhar, F. M., Farooq, J., Saleem, M.
Science, 10, 6465-6468. F., Saeed, A., Nazeer, W., Riaz, M., Naeem, M., &
Ahuja, S. L., & Dhayal, L. S. (2007). Combining ability Javaid, A. (2012). Combining ability analysis for within-
estimates for yield and fibre quality traits in 4 ×13 boll yield components in upland cotton (Gossypium
line × tester crosses of Gossypium hirsutum. hirsutum L.). Genetics and Molecular Research, 11,
Euphytica, 153, 87-98. 2790-2800. http://dx.doi.org/10.4238/2012
https://doi.org/10.1007/s10681-006-9244-y Islam, A. A., Era, F. M., Khalequzzaman, M. F., &
Ali, S., Shah, S. H., Ali, G. M., Iqbal, A., Asad Ullah, M., Chakrabarty, S. (2021). Estimation of heterosis in hybrids
& Zafar, Y. (2012). Bt cry toxin expression profile in of upland cotton (Gossypium hirsutum L.) for seed cotton
selected Pakistani cotton genotypes. Current Science, yield and related traits. Journal of Advanced Plant
102(10), 1632-1636. Science, 1, 1-12.
Arif, M., Hussain, N., Yasmeen, A., Naz, S., Anwar, A., Jatoi, W. A., Baloch, M. J., Veesar, N. F., & Panhwar, S. A.
Mushtaq, S., Iqbal, J., Shaheen, A., Aziz, M., (2011). Combining ability estimates from line × tester
Bukhari, S. A. H., & Shah, S. H. (2022). Exogenous analysis for yield and yield components in upland cotton
application of bio-stimulant and growth retardant genotypes. Journal of Agricultural Research, 49, 165-
improved the productivity of cotton cultivars under 172. Retrieved from https://www apply.jar.punjab.gov.pk
different planting arrangement. Brazilian Journal of Kanasagra, J. R., Valu, M., Raval, L. J., & Rupapara, S.
Biology, 82, e238812. https://doi.org/10.1590/1519- (2022). Heterosis, combining ability and gene action for
6984.238812 seed cotton yield and its contributing characters in cotton
Aydin, U., Balci, S., & Cinar, V. M. (2019). Gene action (Gossypium hirsutum L.). Pharma Innovation
and combining ability for neps and seed coat neps in Journal, 11, 2050-2056.
cotton (Gossypium hirsutum L.). Turkish Journal of Karademir, E., Karademir, C., & Başal, H. (2016). Combining
Field Crops, 24, 211-214. ability and line × tester analysis on heat tolerance in
https://doi.org/10.17557/tjfc.568326 cotton (Gossypium hirsutum L.). Indian Journal of
Balci, S., Cinar, V. M., & Unay, A. (2023). Estimating Natural Sciences, 6, 10515-10525.
gene action and combining ability for yield and fiber Keerthivarman, K., Mahalingam, L., Premalatha, N., Boopathi,
quality in cotton (Gossypium hirsutum N.M., Senguttuvan, K., & Manivannan, A. (2022).
L.). Genetika, 55(1), 61-69. Combining ability and heterosis studies in cotton
https://doi.org/10.2298/GENSR2301061B (Gossypium hirsutum L.) for yield and fibre quality
Falconer, D. S., & Mackay, T. F. C. (1996). Introduction to traits. Electronic Journal of Plant Breeding, 13(2), 697-
quantitative genetics. Longman Group. 704. http://dx.doi.org/10.37992/2022.1302.090
Farooq, A., Shakeel, A., Saeed, A., Farooq, J., Rizwan, M., Kempthorne, O. (1957). An introduction to genetic statistics.
Chattha, W. S., Sarwar, Y., & Ramzan, G. (2022). John Wiley and Sons.
Genetic variability predicting breeding potential of Khan, M., Mohibullah, S. N. M., Shah, M. A., Amin, S., &
upland cotton (Gossypium hirsutum L.) for high Asmat, S. (2018). Genetic variability and morphological
temperature tolerance. Research Square, 1, 1-23. characterization of the elite upland cotton germplasm
https://doi.org/10.1186/s42397-023-00144-z (Gossypium hirsutum L.). Journal of Genetics, Genomics
Fatima, A., Saeed, A., Ullah, M. I., Shah, S. A. H., Ijaz, and Plant Breeding, 2, 1-8.
M., Anwar, M. R., ... & Amjad, I. (2022). Estimation Khan, S. U., Ali, S., Shah, S. H., Zia, M. A., Shoukat, S.,
of gene action for the selection of superior parents Hussain, Z., & Shehzad, A. (2023). Impact of nitrogen

19
Muhammad Ahmad et al Journal of Pure and Applied Agriculture (2023) 8(3): 10-21
and phosphorus fertilizers on Cry1Ac protein Nizamani, M. J. B., Baloach, A. W., Buriro, M., Nizamani, G.
contents in transgenic cotton. Brazilian Journal of S., Nizamani, M. R., Nizamani, I. A., & Baloach, G. H.
Biology, 83, e246436. https://doi.org/10.1590/1519- (2017). Genetic distance, heritability and correlation
6984.246436 analysis for yield and fibre quality traits in upland cotton
Khokhar, E. S., Shakeel, A., Maqbool, M. K., Abuzar, M. genotypes. Pakistan Journal of Biotechnology, 14, 29-36.
K., Zareen, S. S., Aamir, M., & Asadullah. (2018). Punjab Agriculture Department [PAD]. (2022). Cotton
Studying combining ability and heterosis in different production technology 2022-23.
cotton (Gossypium hirsutum L.) genotypes for yield Department of Government of Punjab, Pakistan.
and yield contributing traits. Pakistan Journal of Retrieved from
Agricultural Research, 31, 55-68. https://www.agripunjab.gov.pk/publication
http://dx.doi.org/10.17582/journal.pjar/2018/31.1.55. Rani, S., Chapara, M., & Satish, Y. (2020). Heterosis for seed
68 cotton yield and yield contributing traits cotton
Kumar, A., Nirania, K. S., & Bankar, A. H. (2017). (Gossypium hirsutum L.). International journal of
Combining ability for seed cotton yield and chemical studies, 8, 2496-2500.
attributing traits in American cotton (Gossypium https://doi.org/10.22271/chemi.2020.v8.i3aj.9581
hirsutum L.). Journal of Pharmacognosy and Riaz, A., Ullah, F., Chohan, S., Saeed, A., Aqeel, M., Sher, A.,
Phytochemistry, 6, 376-378. ... & Khalid, M. (2023). Estimation of heterosis and
Makhdoom, K., Khan, N. U., Khan, S. U., Gul, Z., Bibi, combining ability effects for yield and fiber quality traits
R., Gul, S., Ali, N., Ali, S. M., & Khan, N. (2019). in cotton (Gossypium hirsutum L.). Biological and
Genetic effects assessment through line × tester Clinical Sciences Research Journal, 1, 261-261.
combining ability for development of promising https://doi.org/10.54112/bcsrj.v2023i1.261
hybrids based on quantitative traits in Gossypium Sajjad, M., Azhar, M. T., & Malook, M. U. (2016). Line ×
hirsutum L. Journal of Agricultural Sciences, 25, 47- tester analysis for different yield and its attributed traits in
61. https://doi.org/10.15832/ankutbd.538997 upland cotton (Gossypium hirsutum L.). Agricultural
Manan, A., Zafar, M. M., Ren, M., Khurshid, M., Sahar, Biology Journal of North America, 7, 163-172.
A., Rehman, A., ... & Shakeel, A. (2022). Genetic https://doi.org/10.5251/abjna.2016.7.4.163.172
analysis of biochemical, fiber yield and quality traits Salman, M., Zia, Z. U., Iqbal, R. A., Maqsood, R. H., Ahmad,
of upland cotton under high-temperature. Plant S., Bakhsh, A., & Azhar, T. M. (2019). Genetic effects
Production Science, 25(1), 105-119. conferring heat tolerance in upland cotton (Gossypium
https://doi.org/10.1080/1343943X.2021.1972013 hirsutum L.). Journal of Cotton Research, 2, 1-8.
Mawblei, C., Premalatha, N., Rajeswari, S., & https://doi.org/10.1186/s42397-019-0025-2
Manivannan, A. (2022). Genetic variability, Shaukat, M., Ahmad, A., Khaliq, T., Afzal, I., Muhammad, S.,
correlation and path analysis of upland cotton Safdar, B., & Shah, S. H. (2021). Foliar spray of natural
(Gossypium hirsutum L.) germplasm for seed cotton and synthetic plant growth promoters accelerates growth
yield. Electronic Journal of Plant Breeding, 13(3), and yield of cotton by modulating photosynthetic
820-825. https://doi.org/10.37992/2022.1303.104 pigments. International Journal of Plant Production, 15,
Max, S. M., Gibely, H. R., & Abdelmoghny, A. M. (2021). 615-624.
Combining ability in relation to heterosis effects and Shoukat, S., Anwar, M., Ali, S., Begum, S., Aqeel, M., Zia, M.
genetic diversity in cotton using line x tester mating A., Shah, S. H., & Ali, G. M. (2020). Estimation of
design. Plant Archive, 21(1), 1-9. morphological and molecular diversity of seventy-two
Monicashree, C., Amala Balu, P., & Gunasekaran, M. advanced Pakistani cotton genotypes using simple
(2017). Heterosis studies for yield and fibre quality sequence repeats. Applied Ecology and Environmental
traits in upland cotton (Gossypium hirsutum Research, 18(2), 2043-2056.
L.). International Journal of Pure and Applied Shuli, F., Jarwar, A. H., Wang, X., Wang, L., & Ma, Q. (2018).
Biosciences, 5(3), 169-186. Overview of the cotton in Pakistan and its future
http://dx.doi.org/10.18782/2320-7051.2934 prospects. Pakistan Journal of Agricultural Sciences, 31,
Munir, S., Qureshi, M. K., Shahzad, A. N., Manzoor, H., 396-407.
Shahzad K., Aslam, M. (2018). Assessment of gene https://doi.org/10.17582/journal.pjar/2018/31.4.396.407
action and combining ability for fibre and yield Singh, P. K., Singh, K. K., Yadav, B., Kumar, A., &
contributing traits in interspecific and intraspecific Srivastava, A. B. (2022). Constraints in the Production
hybrids of cotton. Czech Journal of Genetics and and Marketing of Rose and Marigold. The Pharma
Plant Breeding, 54, 71-77. Innovation Journal, 11(6), 1463-1466.
https://doi.org/10.17221/54/2017-CJGPB Sivia, S. S., Siwach, S. S., Sangwan, O., Lingaraja, L., &
Nagarajan, S., Purushothaman, Y., Selvavinayagam, M., Vekariya, R. D. (2017). Combining ability estimates for
Govindharaj, P., & Musthafa, A. (2022). Studies on yield traits in line × tester crosses of upland cotton
Colored Cotton: Biochemical and Genetic Aspects. (Gossypium hirsutum). International Journal of Pure and
In Cotton. IntechOpen. Applied Bioscience, 5, 464-474.
https://doi.org/10.5772/intechopen.104898

20
Muhammad Ahmad et al Journal of Pure and Applied Agriculture (2023) 8(3): 10-21
Soomro, A. W. (2020). Estimation of genetic variability Zafar, M. M., Razzaq, A., Farooq, M. A., Rehman, H. Ur.,
parameters in segregating F2 generation of cotton. Firdous, A., Shakeel, A., Mo, M., Ren, M., Ashraf, Y.,
FUUAST Journal of Biology, 10, 83-87. Youlu, Y. (2022). Genetic variation studies of ionic and
Steel, R. G. D., Tome, J. H., & Dickey, D. A. (1997). within boll yield components in cotton (Gossypium
Principle and procedure of statistics: A biometrical hirsutum L.) under salt stress. Journal of Natural Fibers,
approach (3rd ed.). McGraw-Hill. 19, 3063-3082.
Tantuway, G., & Patil, S. S. (2020). L×T analysis in F5 https://doi.org/10.1080/15440478.2020.1838996
lines of RGR population derived through the Zapadiya, V. J., & Valu, M. G. (2021). Gene action studies in
exploitation of heterotic groups in Gossypium the inheritance of yield and quality attributing traits in
species. Journal of Applied Science and Technology diallel cross of cotton (Gossypium hirsutum L.).
Trends, 1, 115-126. International Journal of Pure & Applied Bioscience, 9(3),
https://doi.org/10.9734/CJAST/2020/v39i1730759 105-109. http://dx.doi.org/10.18782/2582-2845.8713
Thiyagu, K., Gnanasekaran, M., & Gunasekaran, M. Zhang, J. F., Abdelraheem, A., & Wu, J. X. (2017). Heterosis,
(2019). Combining ability and heterosis for seed combining ability and genetic effect, and relationship
cotton yield, its components and fibre quality traits in with genetic distance based on a diallel of hybrids from
upland cotton (Gossypium hirsutum L.). Electronic five diverse Gossypium barbadense cotton genotypes.
Journal of Plant Breeding, 10, 1501-1511. Euphytica, 213, 1-13.
https://doi.org/10.5958/0975-928X.2019.00193.5 Zia, M. A., Jan, S. A., Shinwari, Z. K., Shah, S. H., & Khalil,
Usharani, C. V., Manjula, S. M., & Patil, S. S. (2016). A. T. (2015). Impact of Bt cotton on animal health: A
Estimating combining ability through line × tester review. Global Veterinaria, 14(3), 377-381.
analysis in upland cotton. Research in Environment Zia, M. A., Shah, S. H., Shoukat, S., Hussain, Z., Khan, S. U.,
and Life Sciences, 9, 628-633. Retrieved from & Shafqat, N. (2022). Physicochemical features,
http://rels.comxa.com functional characteristics, and health benefits of
Vadodariya, J.M., Patel, B.C., Patel, M.P., Patel, S.K., & cottonseed oil: A review. Brazilian Journal of Biology,
Panchal, S.D. (2022). Heterosis estimation for seed 82, e243511. https://doi.org/10.1590/1519-6984.243511
cotton yield and its component traits in interspecific Zia, M. A., Shinwari, Z. K., Ali, S., Shah, S. H., Anwar, M., &
hybrids of cotton. Retrieved from Ali, G. M. (2018a). Comparison among different cotton
https://www.thepharmajournal.com (Gossypium hirsutum L.) genotypes with respect to
Wang, H., & Memon, H. (2020). Cotton science and morphological, fibre quality attributes and expression
processing technology. Physical Structure, Properties analysis of Cry1Ac gene. Journal of Animal and Plant
and Quality of Cotton, 5, 79-98. Retrieved from Sciences, 28(3), 819-829.
https://link.springer.com/ Zia, M. A., Shinwari, Z. K., Ali, S., Shah, S. H., Anwar, M., &
Yehia, W. M. B., & El-Hashash, E. F. (2019). Combining Ali, G. M. (2018b). Comparison among different cotton
ability effects and heterosis estimates through line × (Gossypium hirsutum L.) genotypes with respect to
tester analysis for yield, yield components and fiber morphological, fibre quality attributes and expression
traits in Egyptian cotton. Journal of Agronomy, 18, analysis of Cry1Ac gene. Journal of Animal and Plant
53238-53246. Sciences, 28(3), 819-829.

21