International Journal of Plant Breeding and Crop Science

Vol. 7(2), pp. 824-832, September, 2020. © www.premierpublishers.org, ISSN: 2167-0449

Research Article

Combining Ability Analysis of Maize (Zea Mays L.) Inbred

Lines for Grain Yield and Yield Related Traits
*Mamud Aman Tello1 and Habtamu Zelleke2
1Maize Breeder, Wondo Genet Agricultural Research Center, Hawassa Maize, Ethiopian Institute of Agricultural Research,
Addis Ababa, Ethiopia
2School of Plant Science, Haramaya University, Haramaya, Ethiopia

A total of 64 test-crosses generated by crossing 32 elite maize inbred lines with two testers and
two standard checks were evaluated for grain yield and yield related traits in 6×11 alpha lattice
design replicated twice during 2017 cropping season at Bako National Maize Research Center of
Ethiopia with the objective of estimating general and specific combining ability effects of the
inbred lines for grain yield and yield related traits. Analysis of variance indicated highly significant
mean squares due to genotypes for all the studied traits. Mean squares due to line general
combining ability (GCA) were significant for all studied traits whereas, mean square due to tester
GCA was significant for all traits, except number of kernels per row and grain yield. Mean squares
due to specific combining ability (SCA) effects were significant (P<0.01 or P<0.05) for biomass
yield, number of ears per plant and thousand kernel weight. Generally, mean squares due to both
lines and testers GCA and SCA of line × tester interactions were significant for grain yield and
most yield related traits indicating the importance of both additive and non-additive gene actions
in controlling these traits.

Key words: line × tester, combining ability, general combining ability, specific combining ability.

INTRODUCTION

In Ethiopia, maize is one of the most important cereal
crops grown, mainly for food security. Today, it is second
to tef in terms of area but first in terms of productivity
among all cereals in the country (CSA, 2015).It grows from
moisture stress areas to high rainfall areas and from
lowlands to the highlands (Kebede et al., 1993).Despite
the large area under maize, the national average yield of
maize is about 3.9 t/ha (CSA, 2018). This is by far below
the world’s average yield which is about 5.4 t/ha (FAO,
2016). Unavailability of suitable maize varieties is one of
the possible reasons responsible for such a yield gap.
Efforts are, therefore, required to be made to develop high
yield potential with biotic and abiotic stress tolerant maize
hybrids in order to increase maize production and fill this
gap.
program that needs to be tested in hybrid combinations in
order to gather information on the combining ability of the
newly developed inbred lines.
This effort is aimed at identifying better combining inbred
lines for more productive, biotic and abiotic stress tolerant
hybrids for mid-altitude areas of the country that can
replace the previously released varieties. Therefore, this
study was initiated to generate information on 32 mid-
altitude elite maize inbred lines crossed to two testers of
known heterotic groups using line × tester mating design
with the objective of estimation of the GCA and SCA
effects of the inbred lines for grain yield and yield related
traits.

Bako National Maize Research Program (BNMRP) in
collaboration with International Maize and Wheat
Improvement Center (CIMMYT) aimed has developed a
large number of medium and early maturing elite maize
inbred lines. As an integral component of the program, new
inbred lines are being continuously developed by the
*Corresponding Author: Mamud Aman Tello, Maize
Breeder, Wondo Genet Agricultural Research Center,
Hawassa Maize, Ethiopian Institute of Agricultural
Research, Addis Ababa, Ethiopia.
E-mail: mamudaman@gmail.com; Co-Author 2Email:
wubeno@gmail.com

Combining Ability Analysis of Maize (Zea Mays L.) Inbred Lines for Grain Yield and Yield Related Traits
Tello and Zelleke. 825

MATERIALS AND METHODS 19.7oC, respectively. Sixty percent of the soil of Bako
National Maize Research Center is reddish brown in colour
Experimental Site with clay and loam texture.
The experiment was conducted at Bako National Maize
Research Center (BNMRC) during the 2017 main cropping The Experimental Materials
season. The area is located in East Wollega Zone of the
Sixty-four test-crosses produced by crossing 32 elite
Oromia Regional State, Western Ethiopia at about 250 km
inbred lines to two testers (PO'00E-3-2-1-2-1-#-# and
west of Addis Ababa, the capital city of the country. It lies
ILO'00E-1-9-1-1-1-1-1-#), and two standard checks
between 9o6' North latitude and 37o09' East longitude in
(BH546 and BH547) were used for the study. Tester,
the sub-humid agro-ecology, with an altitude of 1650 m
PO'00E-3-2-1-2-1-#-# belongs to heterotic group A and
above sea level. The mean annual rainfall of the study area
ILO'00E-1-9-1-1-1-1-1-# has been identified as heterotic
during the study season was 1317mm.The climate of the
group B. All the lines and testers were developed at
area is characterized as sub-humid with mean monthly
BNMRC, and the standard checks were released by the
maximum and minimum temperature of 24.5oC and
same center.
Table 1. Descriptions of the lines, testers and checks used for the study at Bako during 2017
Line code Pedigree Line code Pedigree
L1 S719(F2)-4-2-1-1 L20 P12W(F2)-162-1-1-1
L2 S719(F2)-25-1-1-1 L21 P12W(F2)-162-1-1-2
L3 P63(F2)-27-2-1-1 L22 P12W(F2)-194-1-1-2
L4 P63(F2)-31-1-1-1 L23 P12W(F2)-207-1-1-1
L5 P63(F2)-31-1-1-2 L24 SY312/442(F2)-36-2-1-1
L6 P12W(F2)-4-1-1-1 L25 SY312/442(F2)-84-2-1-1
L7 P12W(F2)-14-1-1-1 L26 SY312/442(F2)-231-1-1-1
L8 P12W(F2)-24-1-1-1 L27 SY312/442(F2)-234-2-1-1
L9 P12W(F2)-24-2-1-1 L28 SY395/202(F2)-28-1-1-1
L10 P12W(F2)-24-2-2-1 L29 SY395/202(F2)-66-1-1-1
L11 P12W(F2)-34-1-1-2 L30 P8M-91(F2)-12-1-1-1-1
L12 P12W(F2)-80-1-2-1 L31 P8M-91(F2)-37-2-1-1-1
L13 P12W(F2)-82-1-1-1 L32 P8M-91(F2)-37-2-2-1-1
L14 P12W(F2)-95-1-1-1 Testers
L15 P12W(F2)-95-2-1-1 T1 ILO'00E-1-9-1-1-1-1-1-# (B)
L16 P12W(F2)-96-1-2-1 T2 PO'00E-3-2-1-2-1-#-# (A)
L17 P12W(F2)-141-2-1-1 Standard Checks
L18 P12W(F2)-161-1-2-1 Check 1 BH546
L19 P12W(F2)-161-1-2-2 Check 2 BH547
Experimental Design and Field Management from planting to when 50% of the plants in aplot produced
2-3 cm long silk. Days to maturity (MD): Number of days
The experiment was conducted in a 6×11 alpha-lattice from planting to when 50% of the kernels on the cob
design with two replications. Each of the 11 incomplete showed black layer on the tip the kernel where it attaches
blocks in replication was comprised of 6 plots having to the cob. Biomass yield (BY): Total aboveground
5.1meter row length with the spacing of 75 cm between biomass yield after oven drying to a constant weight in tons
rows and 30 cm between plants. First, two seeds were per hectare obtained from each plot at harvest. Harvest
planted per hill and later thinned out to one plant per hill index (HI): The ratio of dried grain weight per ha adjusted
after the seedlings established well to give a final plant to 12.5% moisture content to the dried total aboveground
population of 44,444 plants per ha. Fertilizer was applied biomass weight per ha. Number of ears per plant (EPP):
at the rates of 100 kg/ha P2O5 and 100 kg/ha N as Di- Total number of harvested ears in each plot divided by the
ammonium phosphate (DAP) and urea respectively. All stand count at harvest. Number of kernel rows per ear
other cultural practices such as land preparation, weeding, (KRPE): Total numbers of kernel rows of the ear were
disease and insect pest control were done manually as counted from five randomly taken ears and the average
per required. value was used as kernel rows per ear. Number of kernels
per row (KPR): Number of kernels in each row from five
Data Collected randomly taken ears and the average values were
recorded as kernels per row. Thousand kernel weight
Grain yield (GY): The total grain yield in kg per plot and (TKW): 1000 randomly taken kernels were weighed from
adjusted to 12.5% moisture level. Days to anthesis (AD): each plot using sensitive balance and was adjusted to
Number of days from planting to when 50% of the plant in 12.5% moisture level.
a plot shed pollen. Days to silking (SD): Number of days
Combining Ability Analysis of Maize (Zea Mays L.) Inbred Lines for Grain Yield and Yield Related Traits

Data Analysis
The data collected for all yield and yield related traits were
analyzed using PROC MIXED procedure in SAS (SAS,
2004). Traits that showed significant differences were
further subjected to Duncan’s Multiple Range Test (DMRT)
for mean separation. In the analysis, entries were used as
a fixed factor while replications and incomplete blocks
within replication were considered as random factors.
Entry means adjusted for block effects as per the lattice
design (Cochran and Cox, 1957) and it was used for line×
tester analysis to partition the sum of squares due to
crosses into line, tester and line by tester effects
(Dabholkar, 1999, Singh and Chaudhary, 1985) using SAS
computer program (SAS, 2004) and AGD-R software for
traits that showed significant differences among crosses.
The significant of GCA and SCA effects were tested by
dividing the corresponding GCA and SCA values by their
respective standard error and comparing the obtained t
with tabular t-value at error degree of freedom. Square root
transformation was used for normalize root and stem
lodging as suggested by Gomez and Gomez (1984),
whereas𝑙𝑛√(𝐴𝑆𝐼 + 10)was used for normalize anthesis-
silking interval (ASI) as suggested by Bolanos and
Edmeades (1996).
RESULTS AND DISCUSSION
Analysis of Variance (ANOVA)
Analysis of variance was conducted for all the traits and it
showed that mean squares due to genotypes were
significant (P<0.01 or P<0.05) for all traits. The significant
mean squares due to genotypes for all of the traits studied
indicated the existence of variability among the genotypes,
which could be exploited for the future improvement of the
traits. The current finding is in line with the findings of
Shushay et al. (2013).
Mean Performance of Genotypes
The highest grain yield was 11.24 t/ha for the hybrid
L17×T1 followed by L10×T2 (10.87 t/ha), L3×T2
(10.62t/ha), L14×T1 (10.53 t/ha) and L10×T1 (10.44 t/ha),
all of which significantly out yielded the high yielding check,
BH547 (7.96t/ha).Crosses those had better mean values
than the standard checks, indicating the possibility of
obtaining good hybrid combinations for further breeding
and commercialization in the future breeding program.
Hybrids L24×T2 were the earliest, which required only 65
days to anthesis and days to silking respectively. Maturity
days ranged from 130 days for L24×T2 to154 days for
L32×T1.Furthermore,mean value of biomass yield,
harvest index, number of ears per plant and thousand
kernel weight were in the range of54.2 t/h for L32×T2 to
14.5t/h forL29×T2, 60.6 % for L10×T2 to 25.1 %
forL32×T2, 1.6 earsfor L3×T2to 0.9ears for L4×T1, and
531.0 g for L2×T2 to 302.0 g for L3×T1, respectively (Table
Combining Ability Analysis of Maize (Zea Mays L.) Inbred Lines for Grain Yield and Yield Related Traits
Int. J. Plant Breed. Crop Sci. 826
2). The finding indicated that the crosses were sufficiently
different from each other for these traits; and hence,
selection is possible to identify the most desirable cross
combinations.
Combining Ability Analysis
According to Line × tester analysis for grain yield and yield
related traits, GCA mean squares due to lines were highly
significant (P<0.01) for grain yield, biomass yield, 1000
kernel weight, number of rows per ear, number of ears per
plant, number of kernels per row, harvest index, days to
anthesis, silking and maturity. Mean squares for GCA due
to testers were also highly significant for biomass yield,
1000 kernel weight, number of rows per ear, harvest index,
days to anthesis, silking and maturity, and significant
(P<0.05) for number of ears per plant.
Analysis of variance for SCA also showed highly
significant differences for biomass yield, 1000 kernel
weight and number of ears per plant. These indicated that
both additive and non-additive gene effects were involved
in the control of most of the traits. However, the proportion
of GCA sum of squares was higher than that of SCA for all
traits, except for biomass yield, number of ears per plant,
grain yield and thousand kernel weights. This showed the
greater contribution of the additive gene effects than the
non-additive to the genetic variability of the traits in the
crosses (Table 3).
Estimates of General Combining Ability Effects
Estimates of GCA effects for grain yield showed that out of
the 32 inbred lines studied in line × tester mating design,
only L10 exhibited positive and highly significant GCA
effects while L4, L29 and L32 exhibited negative and
significant (P<0.01or P<0.05) GCA effects. Inbred line L10
was a good general combiner for grain yield with the
maximum GCAeffect of (2.32 t/ha), while L32 exhibited the
lowest GCA effect of (-2.20 t/ha) (Table 4). Inbred lines,
L10 also showed highly significant and positive GCA
effects for harvest index.
Both negative and positive significant (P<0.01 or P<0.05)
GCA effects for days to anthesis and maturity were
exhibited. Accordingly, L4, L19 and L24 were exhibited
negative and highly significant GCA effects for days to
anthesis and days to maturity while L2, L30 and L32 were
exhibited positive and highly significant GCA effects for
both days to anthesis and days to maturity, respectively.
Inbred line L4 and L 24 exhibited negative and significant
(P<0.01or P<0.05) GCA effects for days to silking. This
indicated that the possibility of making an effective
selection for these traits to develop early maturing hybrids.
From the tester, T2 showed negative and highly significant
GCA effects whereas T1 showed positive and highly
significant GCA effects for days to anthesis, silking and
maturity. Both positive and negative GCA effects for days
to anthesis and maturity were reported by Gudeta (2007)
and Shengu et al. (2016).
Tello and Zelleke. 827

Table 2. Mean of grain yield and yield related traits of selected F1maize hybrids and two standard checks
Code GY DA DS DM BY HI EPP RPE KPR TKW
L1×T1 8.445 72.0 72.0 144.0 23.15 36.60 1.2 14 36.0 411.5
L1×T2 8.625 68.5 69.00 137.0 22.35 38.55 1.0 14 41.0 451.5
L2×T1 8.360 73.5 74.00 147.0 21.95 38.15 1.1 14 36.0 395.0
L2×T2 9.805 72.0 73.00 144.0 24.35 40.25 1.2 14 40.5 531.0
L3×T1 6.975 70.5 71.00 141.0 19.00 36.60 1.3 14 40.0 302.0
L3×T2 10.620 65.0 66.00 130.0 26.30 39.60 1.6 13 35.5 391.5
L4×T1 5.605 67.0 68.00 134.0 17.70 32.10 0.9 14 41.0 397.0
L4×T2 7.205 66.5 68.00 133.0 16.45 43.80 0.9 13 41.5 412.5
L5×T1 8.045 71.5 72.00 143.0 22.00 36.35 0.9 14 39.5 409.0
L5×T2 9.755 68.5 69.50 137.0 22.00 44.35 1.3 14 40.5 410.5
L6×T1 6.540 73.5 74.00 147.0 21.85 30.00 1.0 15 34.5 322.5
L6×T2 8.190 69.0 69.50 138.0 19.80 41.90 1.2 14 40.5 401.0
L7×T1 8.490 70.0 70.00 140.0 25.15 34.20 1.4 15 42.0 356.5
L7×T2 8.575 68.5 69.00 137.0 20.60 41.60 1.2 14 42.0 358.5
L8×T1 9.660 70.5 71.00 141.0 25.90 37.25 1.0 14 42.5 371.0
L8×T2 8.775 68.5 68.50 137.0 25.55 35.00 1.2 14 39.0 380.0
L9×T1 7.865 70.5 70.50 141.0 25.80 30.65 1.1 16 38.5 387.0
L9×T2 8.835 67.0 67.50 134.0 20.35 43.45 0.9 14 42.5 406.0
L10×T1 10.435 69.0 69.00 138.0 26.15 40.00 1.4 16 40.5 345.0
L10×T2 10.870 67.5 67.50 135.0 18.40 60.60 1.0 14 41.5 402.5
L11×T1 9.095 70.5 70.50 141.0 22.55 40.60 1.0 15 41.0 414.5
L11×T2 9.610 69.0 69.50 138.0 23.60 40.85 1.2 15 40.0 438.0
L12×T1 9.345 69.5 70.00 139.0 26.20 35.85 1.4 15 39.5 319.0
L12×T2 7.975 67.5 68.00 135.0 19.45 40.80 1.2 14 42.0 349.5
L13×T1 8.915 67.5 68.00 135.0 23.70 37.50 1.4 15 34.5 371.5
L13×T2 7.950 67.0 68.00 134.0 18.45 43.40 1.0 14 39.0 431.5
L14×T1 10.525 68.5 68.50 137.0 24.10 43.35 1.4 14 39.5 413.5
L14×T2 8.380 69.0 69.50 138.0 19.15 43.65 1.3 14 39.5 341.5
L15×T1 8.915 69.0 69.00 138.0 27.80 32.10 1.1 14 43.5 389.0
L15×T2 6.665 66.5 66.50 133.0 19.50 35.50 1.1 14 42.5 399.5
L16×T1 8.700 71.0 71.00 142.0 22.10 39.35 1.3 15 38.0 360.5
L16×T2 8.310 65.0 66.00 130.0 18.65 44.55 1.1 14 42.0 425.5
L17×T1 11.235 70.0 71.00 140.0 30.85 36.40 1.5 14 39.5 361.0
L17×T2 7.290 67.0 68.00 134.0 21.00 35.90 1.4 14 41.0 373.0
L18×T1 8.755 68.5 69.00 137.0 21.70 40.45 1.1 16 41.0 319.0
L18×T2 8.955 66.5 68.50 133.0 20.65 43.20 1.0 15 41.5 418.0
L19×T1 9.020 67.5 69.00 135.0 24.65 36.65 1.2 18 49.5 336.0
L19×T2 9.870 66.5 69.00 133.0 23.70 41.65 1.1 15 42.5 372.5
L20×T1 10.245 71.5 72.00 143.0 29.35 34.40 1.5 15 42.5 356.5
L20×T2 8.590 70.0 71.00 140.0 20.45 42.10 1.1 14 43.5 350.5
L21×T1 8.600 73.0 73.00 146.0 23.80 36.30 1.3 16 42.0 364.5
L21×T2 6.995 69.5 71.00 139.0 24.35 29.70 1.3 15 40.0 349.0
L22×T1 8.690 72.5 73.00 145.0 23.75 36.75 1.2 14 36.5 401.0
L22×T2 7.710 66.5 68.00 133.0 20.15 38.30 1.2 14 41.0 334.0
L23×T1 9.245 70.0 71.00 140.0 23.75 39.20 1.1 16 45.0 358.5
L23×T2 8.665 66.0 67.00 132.0 23.00 37.65 0.9 15 42.0 381.5
L24×T1 7.450 67.5 67.50 135.0 18.35 40.90 1.3 15 40.0 308.0
L24×T2 7.065 65.0 65.00 130.0 16.30 43.40 1.0 14 40.5 428.0
L25×T1 7.785 70.0 71.00 140.0 22.05 35.25 1.1 15 43.0 358.0
L25×T2 8.570 67.5 68.50 135.0 20.55 41.70 1.1 14 41.5 430.5
L26×T1 7.340 70.0 71.00 140.0 20.80 35.40 1.1 16 39.5 347.0
L26×T2 6.990 67.0 68.00 134.0 15.75 44.25 1.1 14 38.5 320.5
L27×T1 9.670 71.5 72.00 143.0 23.70 40.95 1.0 15 43.0 334.5
L27×T2 7.515 68.0 69.50 136.0 18.60 40.35 1.1 14 40.0 353.5
L28×T1 9.355 72.0 72.50 144.0 23.05 40.60 1.3 14 38.0 348.5
L28×T2 7.215 68.0 68.00 136.0 19.85 36.00 1.2 14 43.0 402.0
L29×T1 7.130 70.0 71.00 140.0 14.60 48.75 1.2 14 38.0 323.0
Combining Ability Analysis of Maize (Zea Mays L.) Inbred Lines for Grain Yield and Yield Related Traits
 Int. J. Plant Breed. Crop Sci. 828
Table 2 Continue. Mean of grain yield and yield related traits of selected F 1maize hybrids and two standard checks
Code GY DA DS DM BY HI EPP RPE KPR TKW
L29×T2 7.075 68.0 69.00 136.0 14.45 49.00 1.1 14 36.0 392.5
L30×T1 8.265 73.5 73.50 147.0 23.20 35.65 1.0 14 42.0 436.5
L30×T2 8.965 72.5 74.50 145.0 24.10 37.10 1.1 14 40.0 437.0
L31×T1 7.230 71.0 71.00 142.0 20.70 34.90 1.1 14 39.0 340.5
L31×T2 8.525 66.5 67.50 133.0 20.95 40.85 1.1 15 38.0 470.0
L32×T1 6.740 77.0 77.50 154.0 26.30 25.95 1.3 14 31.5 388.0
L32×T2 5.980 74.0 75.50 148.0 54.20 25.10 1.2 12 27.5 519.5
Check1 6.946 69.5 71.5 139.0 25.90 28.13 1.0 15 42.5 366.7
Check2 7.959 71.0 71.0 142.0 20.28 39.22 1.1 17 40.1 366.7
Mean 8.419 69.3 69.93 138.5 22.50 38.80 1.2 14 40.4 382.93
GY = Grain Yield, DA = Days to Anthesis, DS = Days to Silking, DM= Days to Maturity, BY = biomass yield, HI = Harvest
Index, EPP = Number of Ears per Plant, RPE = Number of Kernel Rows per Ear, KPR = Number of Kernels per Row,
TKW = Thousand Kernel Weight.

Table 3. Mean squares for combining ability analyses for grain yield and yield related traits evaluated at Bako in 2017
Source of
df GY DA DS DM BY HI EPP RPE KPR TKW
variation
Crosses 63 2.653* 10.687** 12.487** 42.747** 52.249** 77.410** 0.048** 1.579** 43.847* 3520.82**
Line (gca) 31 3.077** 12.831** 16.470** 51.325** 62.812** 104.025** 0.057** 1.919** 57.615** 3918.28**
Tester
1 1.005ns 205.727** 102.560** 822.907** 81.674** 384.684** 0.083* 18.000** 2.114ns 36886.86**
(gca)
L×T(sca) 31 2.283ns 2.251ns 5.598ns 9.003ns 40.737** 40.883ns 0.037** 0.710ns 31.425ns 2047.03**
Error 43 1.543 1.761 5.547 7.042 12.147 25.441 0.016 0.776 26.822 914.65
CV % 14.57 1.89 3.34 1.89 15.34 13.65 11.70 6.60 12.53 7.78
ns, * and **: Non significant, significant at 5% and 1% levels of probability, respectively.
GY = Grain Yield, DA = Days to Anthesis, DS = Days to Silking, DM= Days to Maturity, BY = biomass yield, HI = Harvest
Index, EPP = Number of Ears per Plant, RPE = Number of Kernel Rows per Ear, KPR = Number of Kernels per Row,
TKW = Thousand Kernel Weight.

With respect to harvest index, inbred lines L10 and L29
were good general combiners while inbred line 32 was
poor general combiner for this trait. This indicated that
these inbred lines (L10 and L29) highly contributed to the
partitioning of total biomass to grain, thereby increase
grain yield. Similarly, Shushay et al. (2013) reported
significant GCA effects for this trait in both directions. From
the tester, T2 was positively contributed for harvest index
while T1 showed negative and significant GCA effects for
this trait and hence, good and poor general combiner for
this trait respectively. For number of ears per plant,
positive and highly significant GCA effects manifested by
L3 whereas negative and highly significant GCA effects
were exhibited by L4. In maize, positive and significant
GCA effects for number of ears per plant indicates
prolificacy which is a desirable character and increase the
productivity.
With respect to number of rows per ear and number of
kernels per row, L19 showed positive and highly significant
GCA effects while L32 showed negative and highly
significant GCA effects for both traits. For these traits,
positive and significant GCA effects had desirable
character as it is the most important yield component and
highly contributed to increased grain yield. Regarding the
testers, T2 showed positive and highly significant GCA
effects while T1 showed negative and highly significant
GCA effects for number of rows per ear and as a result, T2
was the best and T1 was the poorest general combiner for
this trait.
Combining Ability Analysis of Maize (Zea Mays L.) Inbred Lines for Grain Yield and Yield Related Traits
For biomass yield, L32 and L17 showed positive and
significant (P<0.01or P<0.05) GCA effects while negative
and highly significant GCA effects exhibited by L4 and L29.
This indicated that L32 and L17 were good general
combiners for total biomass yield. Positive and highly
significant GCA effects for thousand kernel weight
exhibited by L1, L2, L30 and L32 whereas, negative and
highly significant GCA effects for thousand kernel weight
exhibited by L12 and L26.This indicated that L1, L2, L30
and L32 were good general combiners for thousand kernel
weight. Regarding the testers, T1 showed positive and
highly significant GCA effects whereas T2 showed
negative and highly significant GCA effects for thousand
kernel weight and as a result, T1 was the best and T2 was
the poorest general combiner for this trait. Inbred lines
identified for good general combining ability could be
utilized in hybrid development programs for improvement
of the traits of interest as these lines have high potential to
transfer desirable gene to their respective crosses.
Estimation of Specific Combining Ability
Both negative and positive and significant to highly
significant estimates of SCA effects were observed among
the crosses (Table 5). Accordingly, Cross L3×T2 and
L17×T1showed positive and highly significant SCA effects
for grain yield whereas, L3×T1 and L17×T2 showed
negative and highly significant SCA effects, indicating that
the crosses were good specific combiners and poor
specific combiners for grain yield respectively. Highly
Tello and Zelleke. 829

Table 4. Estimates of general combining ability effects for grain yield and yield related traits at Bako in 2017
Line GY DA DS DM BY HI EPP RPE KPR TKW
L1 0.141 1.117 -4.740** 2.234 0.290 -1.007 -0.046 -0.438 -2.717 45.614**
L2 0.488 3.297** 3.693** 6.595** 0.489 0.618 0.004 -0.438 -2.038 75.327**
L3 0.407 -1.434* -1.649 -2.867* 0.215 -0.482 0.228** -0.938* -2.654 -42.046*
L4 -2.047** -2.330** -1.792 -4.660** -5.285** -0.632 -0.381** -0.938* 0.703 23.295
L5 0.528 0.669 0.623 1.338 -0.464 1.768 -0.004 -0.438 -0.393 11.600
L6 -1.012 1.866* 1.735 3.731* -1.738 -2.632 -0.022 0.062 -2.819 -26.110
L7 -0.002 0.159 -0.162 0.319 0.448 -0.682 0.076 0.062 1.511 -11.871
L8 0.719 0.344 0.680 0.687 3.174 -2.457 -0.043 -0.438 -0.101 -14.679
L9 -0.072 -0.826 -1.160 -1.652 0.721 -1.532 -0.101 0.563 0.462 16.962
L10 2.321** -1.145 -1.684 -2.290 0.002 11.718** 0.065 0.563 1.399 -8.808
L11 1.185 0.515 0.161 1.029 0.876 2.143 -0.017 0.563 0.203 36.841*
L12 0.361 -1.128 -1.222 -2.255 0.412 -0.257 0.163* 0.063 1.337 -46.904**
L13 -0.085 -1.772* -1.101 -3.544* -1.288 1.868 -0.022 0.063 -4.215 21.251
L14 0.970 -0.649 -0.964 -1.297 -0.843 4.918 0.155* -0.438 -0.804 -7.866
L15 -0.626 -1.464* -1.779 -2.928* 0.978 -4.782 -0.070 -0.438 2.324 12.801
L16 0.047 -1.268 -0.631 -2.537 -2.193 3.368 0.047 0.063 -1.099 7.823
L17 0.900 -0.533 -0.158 -1.067 3.625* -2.432 0.257* -0.438 -0.371 -9.644
L18 0.304 -1.770* -1.127 -3.540* -1.349 3.243 -0.080 1.063* 0.915 -9.296
L19 1.040 -2.272** -0.338 -4.544** 1.783 0.568 -0.027 2.063** 16.353** -23.011
L20 0.895 1.669* 1.664 3.337* 2.255 -0.332 0.105 0.063 2.437 -39.243*
L21 -0.591 1.728* 2.000 3.455* 1.578 -5.582* 0.160* 1.063* 0.708 -29.677
L22 -0.245 0.235 0.654 0.470 -0.591 -1.057 0.034 -0.438 -1.724 -14.730
L23 0.450 -1.274 -1.273 -2.549 0.759 -0.157 -0.162* 1.063* 3.037 -22.236
L24 -0.979 -2.873** -3.366* -5.747** -5.108* 3.568 -0.016 0.063 -0.076 1.972
L25 -0.311 -0.354 0.196 -0.708 -1.120 -0.107 -0.110 0.063 2.670 24.796
L26 -1.037 -0.865 -0.295 -1.730 -4.080* 1.243 -0.015 0.563 -1.351 -49.946**
L27 0.083 0.649 1.024 1.299 -1.362 2.068 -0.136* 0.063 1.343 -32.949*
L28 0.225 0.715 0.860 1.431 -0.776 -0.282 0.140* -0.438 -0.114 -8.493
L29 -1.289* -0.292 0.121 -0.584 -8.115** 10.293** -0.015 -0.438 -3.266 -26.115
L30 0.056 3.738** 3.840** 7.475** 0.929 -2.207 -0.160* -0.438 0.559 52.027**
L31 -0.518 -0.715 -0.737 -1.430 -1.781 -0.707 -0.040 0.063 -1.834 19.930
L32 -2.204** 6.181** 6.712** 12.363** 17.690** -20.057** 0.034 -1.438** -10.988** 67.775**
SE (lines) 0.62 0.66 1.18 1.33 1.74 2.52 0.06 0.44 2.59 15.12
T1 -0.069 1.332** 0.896** 2.663** -0.793 -1.734* -0.022 -0.375** -0.173 17.542**
T2 0.076 -1.337** -0.909** -2.674** 0.801 1.734* 0.022 0.375** 0.135 -17.892**
SE(Tester
0.16 0.17 0.29 0.33 0.44 0.63 0.02 0.11 0.65 3.78
s)
* and **: significant at 5% and 1% levels of probability, respectively.
GY = Grain Yield, DA = Days to Anthesis, DS = Days to Silking, DM= Days to Maturity, BY = biomass yield, HI = Harvest
Index, EPP = Number of Ears per Plant, RPE = Number of Kernel Rows per Ear, KPR = Number of Kernels per Row,
TKW = Thousand Kernel Weight.
significant SCA effects of the crosses indicate significant effects were good specific combiners, while crosses with
deviation from expectations that have been predicted on positive and significant SCA effects were poor specific
the basis of the average performance (GCA effects) of the combiners for days to anthesis and days to maturity. For
parent involved. The crosses with positive and highly number of days to silking, only cross L1×T2 showed
significant SCA effects can be used in maize breeding negative and highly significant SCA effects and, good
program as such crosses contributed to increase in grain specific combiner for this trait. The existence of both
yield potential of maize. Similar to this finding, Dagne et al. negative and positive SCA effects in maize crosses has
(2010), Yoseph et al. (2011) and Shengu et al. (2016) been also reported by (Gudeta, 2007; Shengu et al., 2016;
reported significant SCA effects for grain yield of maize. Murtadha et al., 2018).

With respect to number of days to anthesis and maturity, Positive and highly significant SCA effects for above
negative and significant SCA effects were exhibited by ground biomass exhibited by L32×T2 indicating that the
crosses L3×T1, L14×T2, L16×T1 and L22×T1 whereas lines involved in the combination were good specific
positive and significant SCA effects exhibited by crosses combiner for this trait, whereas negative and highly
L3×T2, L14×T1, L16×T2 and L22×T2 for both traits. significant SCA effects exhibited by L32×T1 indicating that
Accordingly, crosses with negative and significant SCA this cross was poor specific combiner for the trait. For
Combining Ability Analysis of Maize (Zea Mays L.) Inbred Lines for Grain Yield and Yield Related Traits
 Int. J. Plant Breed. Crop Sci. 830

harvest index, positive and highly significant SCA effects
exhibited by L10×T1 and L32×T2 whereas, negative and
highly significant SCA effects exhibited by L10×T2 and
L32×T1. This indicated that the crosses were good specific
combiners and poor specific combiners for this trait
respectively. Positive and negative SCA effects with
significant to highly significant for number of ears per plant
were observed for some crosses. Positive and highly
significant SCA effects for this trait exhibited by L5×T1 and
L13×T2, while negative and highly significant SCA effects
for this trait exhibited by L5×T2 and L13×T1.Crosses with
positive and highly significant SCA effects were good
specific combiners and crosses with negative and highly
significant SCA effects were poor specific combiners for
this trait.

Table 5. Estimates of specific combining ability (SCA) effects of line ×tester for yield and yield related traits evaluated at
Bako
Code GY DA DS DM BY HI EPP RPE KPR TKW
L1×T1 -0.407 -0.384 5.098** -0.767 -0.505 -0.759 0.013 0.375 2.990 -0.645
L1×T2 0.401 0.389 -5.085** 0.778 0.497 0.759 -0.013 -0.375 -2.953 0.996
L2×T1 -0.940 0.612 0.407 1.225 -2.052 -0.684 0.066 0.375 2.449 52.185*
L2×T2 0.933 -0.607 -0.393 -1.215 2.044 0.684 -0.066 -0.375 -2.411 -51.834*
L3×T1 -2.144** -1.435* -1.278 -2.870* -4.765 -0.234 0.203* -0.125 -2.092 19.027
L3×T2 2.138** 1.440* 1.291 2.881* 4.757 0.234 -0.203* 0.125 2.130 -18.676
L4×T1 -0.900 1.098 0.862 2.195 -0.136 4.116 -0.010 -0.125 0.372 -13.574
L4×T2 0.893 -1.092 -0.848 -2.185 0.128 -4.116 0.010 0.125 -0.335 13.924
L5×T1 -0.912 -0.196 -0.010 -0.392 -0.806 2.266 0.249** 0.375 0.489 -22.587
L5×T2 0.905 0.201 0.023 0.402 0.798 -2.266 -0.249** -0.375 -0.451 22.937
L6×T1 -0.828 -0.687 -0.975 -1.374 0.299 4.216 0.084 -0.125 2.924 21.231
L6×T2 0.822 0.692 0.989 1.384 -0.307 -4.216 -0.084 0.125 -2.886 -20.880
L7×T1 0.034 0.534 0.409 1.068 1.588 1.966 -0.074 -0.125 0.079 -19.633
L7×T2 -0.040 -0.529 -0.395 -1.058 -1.596 -1.966 0.074 0.125 -0.042 19.984
L8×T1 0.151 0.454 0.202 0.908 -0.876 -2.859 0.120 0.375 -2.439 -7.777
L8×T2 -0.157 -0.449 -0.188 -0.898 0.868 2.859 -0.120 -0.375 2.477 8.128
L9×T1 -0.601 -0.244 -0.397 -0.488 1.897 4.666 -0.075 -0.625 1.902 0.789
L9×T2 0.594 0.249 0.410 0.499 -1.905 -4.666 0.075 0.625 -1.865 -0.439
L10×T1 0.004 0.540 0.030 1.080 3.220 8.566** -0.200* -0.625 -0.034 13.772
L10×T2 -0.006 -0.535 -0.017 -1.070 -3.228 -8.566** 0.200* 0.625 0.071 -13.421
L11×T1 -0.322 0.490 0.304 0.981 -1.278 -1.609 0.134 0.375 -0.337 -11.766
L11×T2 0.316 -0.485 -0.290 -0.971 1.270 1.609 -0.134 -0.375 0.374 12.116
L12×T1 0.631 0.384 0.019 0.767 2.583 0.741 -0.096 -0.125 1.849 1.052
L12×T2 -0.638 -0.378 -0.005 -0.757 -2.591 -0.741 0.096 0.125 -1.811 -0.702
L13×T1 0.542 1.104 0.438 2.207 2.059 1.216 -0.239** -0.125 2.999 0.658
L13×T2 -0.548 -1.098 -0.424 -2.197 -2.067 -1.216 0.239** 0.125 -2.961 -0.307
L14×T1 1.053 1.626* 1.590 3.253* 1.728 -1.584 -0.041 0.375 0.379 -49.243*
L14×T2 -1.059 -1.621* -1.576 -3.243* -1.736 1.584 0.041 -0.375 -0.342 49.594*
L15×T1 1.006 0.033 -0.527 0.066 3.314 -0.034 0.014 0.375 -0.101 -5.118
L15×T2 -1.012 -0.028 0.540 -0.056 -3.322 0.034 -0.014 -0.375 0.139 5.469
L16×T1 -0.088 -1.472* -1.061 -2.945* 0.670 0.866 -0.058 -0.125 1.606 24.290
L16×T2 0.082 1.477* 1.074 2.955* -0.678 -0.866 0.058 0.125 -1.569 -23.939
L17×T1 1.818** -0.194 -0.512 -0.388 3.994 -1.984 0.031 0.375 0.768 -16.491
L17×T2 -1.825** 0.199 0.525 0.399 -4.002 1.984 -0.031 -0.375 -0.731 16.841
L18×T1 -0.254 0.246 0.370 0.492 -0.175 -0.359 -0.012 -0.125 0.600 21.209
L18×T2 0.247 -0.241 -0.356 -0.481 0.167 0.359 0.012 0.125 -0.562 -20.859
L19×T1 -0.417 0.777 0.348 1.555 -0.158 0.766 -0.036 -1.125** -12.755** -1.003
L19×T2 0.411 -0.772 -0.335 -1.545 0.150 -0.766 0.036 1.125** 12.792** 1.353
L20×T1 0.737 0.693 0.953 1.387 3.688 2.116 -0.164 -0.125 0.877 -17.357
L20×T2 -0.743 -0.688 -0.939 -1.377 -3.696 -2.116 0.164 0.125 -0.840 17.708
L21×T1 0.648 -0.457 -0.205 -0.915 -1.056 -5.034 -0.003 -0.125 -0.711 -20.673
L21×T2 -0.655 0.462 0.219 0.925 1.048 5.034 0.003 0.125 0.748 21.024
L22×T1 0.577 -1.520* -1.573 -3.040* 1.040 -0.959 -0.014 0.375 2.429 -46.097*
L22×T2 -0.583 1.525* 1.586 3.051* -1.048 0.959 0.014 -0.375 -2.392 46.448*
L23×T1 -0.084 -0.656 -0.812 -1.313 -0.711 -2.509 0.013 -0.125 -1.401 -12.073
L23×T2 0.077 0.662 0.825 1.323 0.703 2.509 -0.013 0.125 1.439 12.423
L24×T1 0.054 0.062 -0.418 0.124 0.237 -0.484 -0.159 -0.125 0.515 42.542
Combining Ability Analysis of Maize (Zea Mays L.) Inbred Lines for Grain Yield and Yield Related Traits
Tello and Zelleke. 831

Table 5 Continue. Estimates of specific combining ability (SCA) effects of line ×tester for yield and yield related traits
evaluated at Bako
Code GY DA DS DM BY HI EPP RPE KPR TKW
L24×T2 -0.060 -0.057 0.432 -0.113 -0.245 0.484 0.159 0.125 -0.478 -42.192
L25×T1 -0.350 0.217 -0.181 0.433 0.040 1.491 0.033 -0.125 -1.184 15.071
L25×T2 0.343 -0.211 0.195 -0.423 -0.048 -1.491 -0.033 0.125 1.222 -14.720
L26×T1 -0.134 -0.012 -0.426 -0.023 1.589 2.691 0.060 -0.625 -0.317 -32.512
L26×T2 0.128 0.017 0.439 0.034 -1.597 -2.691 -0.060 0.625 0.355 32.862
L27×T1 0.846 -0.575 -0.596 -1.151 1.631 -2.034 0.057 -0.125 -1.227 -6.483
L27×T2 -0.853 0.580 0.609 1.161 -1.640 2.034 -0.057 0.125 1.264 6.834
L28×T1 0.929 -0.820 -1.974 -1.640 0.683 -4.034 0.004 0.375 3.539 1.142
L28×T2 -0.935 0.825 1.987 1.651 -0.691 4.034 -0.004 -0.375 -3.501 -0.791
L29×T1 0.217 0.172 -0.181 0.344 -0.613 -1.609 0.016 0.375 -0.760 6.851
L29×T2 -0.224 -0.167 0.194 -0.333 0.605 1.609 -0.016 -0.375 0.797 -6.501
L30×T1 -0.679 0.858 1.493 1.716 -1.498 -1.009 0.117 0.375 -0.544 -15.523
L30×T2 0.672 -0.853 -1.480 -1.706 1.490 1.009 -0.117 -0.375 0.582 15.874
L31×T1 -0.629 -1.129 -1.256 -2.258 -1.067 1.241 0.043 0.875* 0.008 41.274
L31×T2 0.623 1.134 1.269 2.269 1.059 -1.241 -0.043 -0.875* 0.030 -40.923
L32×T1 0.344 -0.036 0.075 -0.071 -14.691** -9.159** -0.077 -0.625 -2.272 43.070*
L32×T2 -0.350 0.041 -0.062 0.081 14.683** 9.159** 0.077 0.625 2.310 -42.720*
SE (LxT) 0.88 0.94 1.67 1.88 2.46 3.57 0.09 0.62 3.66 21.39
ns, * and **: Non significant, significant at 5% and 1% levels of probability, respectively.
GY = Grain Yield, DA = Days to Anthesis, DS = Days to Silking, DM= Days to Maturity, BY = biomass yield, HI = Harvest
Index, EPP = Number of Ears per Plant, RPE = Number of Kernel Rows per Ear, KPR = Number of Kernels per Row,
TKW = Thousand Kernel Weight.

For number of rows per ear, crosses L19×T2 and L31×T1
showed positive and highly significant (P<0.01) and
significant (P<0.05) SCA effects respectively. Crosses
L19×T1 and L31×T2 exhibited negative and highly
significant (P<0.01) and significant (P<0.05) SCA effects
respectively for the same trait. The positive SCA effect is
desired for number of rows per ear as it is directly
contributes to increased grain yield and hence, Crosses
L19×T2 and L31×T1 were the best specific combiners for
this particular trait while L19×T1 and L31×T2 were the
worst specific combiners as the crosses were produced
highly significant positive and negative SCA effects,
respectively.
With respect to number of kernels per row, only cross
L19×T2 exhibited positive and highly significant SCA effect
whereas L19×T1 showed negative and highly significant
SCA effects (Table 5).This indicated that most of the
crosses evaluated in the current study did not
significantly deviate from what would have been predicted
based on their parental performance (GCA) effect.
Accordingly, L19×T2 was good specific combiner, while
L19×T1 was poor specific combiner for this trait. For
thousand kernel weight, 8 crosses showed significant
estimates of SCA effects (Table 5). Positive and significant
SCA effects were observed for L2×T1, L14×T2, L22×T2
and L32×T1 while negative and significant SCA effects
were observed for L2×T2, L14×T1, L22×T1 and L32×T2
for this trait. Crosses with positive and significant SCA
effects for this trait are desirable as this trait is the most
important yield component for maize and hence, crosses
with positive and significant SCA effect could be selected
for further use in the breeding program to increase
productivity. The results of the current study are in
agreement with the findings of Shushay et al. (2013),
Habtamu (2015), Shengu et al. (2016) and Murtadha et al.
(2018).
CONCLUSION
Generally, the results of the current study identified that
inbred lines with good GCA, cross combinations with
desirable SCA for the studied traits. Accordingly, Parent
L10 and Cross L3 × T2 exhibited the highest GCA and
SCA for grain yield respectively. This indicates the
possibility of developing desirable hybrids and synthetic
varieties through crossing and or recombination of inbred
lines with desirable traits of interest. In addition, inbred
lines identified for good GCA and the promising crosses
observed for SCA can be utilized in maize breeding
programs for improvement of the traits of interest as well
as for direct release after confirming the stability of their
performances observed in the current study and could be
assisted with a molecular marker-assisted breeding to
identify quantitative trait loci (QTL) responsible for traits of
interest.
ACKNOWLEDGEMENT
The authors are thankful to the Ethiopian Institute of
Agricultural Research (EIAR) for providing the chance to
join the postgraduate study and all the financial support for
this study. Great thanks owed to Haramaya University
for educating me towards my MSc degree.

Combining Ability Analysis of Maize (Zea Mays L.) Inbred Lines for Grain Yield and Yield Related Traits

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Accepted 26 September 2019
Citation: Tello MA, Zelleke H (2020). Combining Ability
Analysis of Maize (Zea Mays L.) Inbred Lines for Grain
Yield and Yield Related Traits. International Journal of
Plant Breeding and Crop Science, 7(2): 824-832.
Copyright: © 2020: Tello and Zelleke. This is an open-
access article distributed under the terms of the Creative
Commons Attribution License, which permits unrestricted
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provided the original author and source are cited.