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Genetic variability and agronomic performances of rice genotypes in different

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Article in Journal of Agriculture and Food Research · August 2023

DOI: 10.1016/j.jafr.2023.100750

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 Journal of Agriculture and Food Research 14 (2023) 100750

Contents lists available at ScienceDirect

Journal of Agriculture and Food Research

journal homepage: www.sciencedirect.com/journal/journal-of-agriculture-and-food-research

Genetic variability and agronomic performances of rice genotypes in

different growing seasons in Bangladesh
Md. Arifur Rahman Khan a, c, 1, *, Apple Mahmud a, 1, Md. Nahidul Islam b, d,
Uttam Kumar Ghosh a, Md. Saddam Hossain a
a
Department of Agronomy, Bangabandhu Sheikh Mujibur Rahman Agricultural University, Gazipur, 1706, Bangladesh
b
Department of Agro-Processing, Bangabandhu Sheikh Mujibur Rahman Agricultural University, Gazipur, 1706, Bangladesh
c
Institute of Genomics for Crop Abiotic Stress Tolerance, Department of Plant and Soil Science, Texas Tech University, Lubbock, TX, 79409, USA
d
Institute of Food Safety and Processing, Bangabandhu Sheikh Mujibur Rahman Agricultural University, Gazipur, 1706, Bangladesh

A R T I C L E I N F O A B S T R A C T

Keywords: Genetic variability is a key factor in the selection of suitable genotypes in rice breeding programs. To evaluate the
Rice genetic variability of 11 rice genotypes, a study was carried out in the field of agronomy at Bangabandhu Sheikh
Agronomic traits Mujibur Rahman Agricultural University (BSMRAU), Gazipur, during 2019–2020. These genotypes were char­
Genetic variations
acterized for 11 traits and analyzed to determine the level of genetic and agronomic diversity as well as the
Heritability
Correlation
degree of association existing between grain yield and its related component traits. The results revealed sig­
nificant differences (P ≤ 0.001 and 0.05) among the rice genotypes for the studied traits in all growing seasons.
Furthermore, it was noted that the phenotypic coefficient of variation was more pronounced than the genotypic
coefficient of variation, highlighting the environmental impact on each trait. The majority of the traits had
moderate to high heritability and genetic advance across the three seasons, showing additive gene action. The
principal component analysis showed that the first three principal components accounted for the greatest
variability, with the majority of the evaluated traits significantly influencing the genetic variability. The mean
value of studied traits exhibited that there was seasonal variation of grain yield by the genotypes, such as BU-R-
ACC-08 and BU-R-ACC-11 genotypes, which produced higher yield in Aus, BU-R-ACC-05 and BU-R-ACC-06 in
Aman, and BU-R-ACC-07 and BU-R-ACC-06 in Boro season, where several genotypes mature earlier in all
growing seasons, viz., BU-R-ACC-05, BU-R-ACC-01, BU-R-ACC-10, and BU-R-ACC-11. The correlation analysis
revealed that there were significant, strong positive or negative correlations among the traits. The results suggest
that agronomical traits that were positively correlated with grain yield could be useful in selecting desired rice
genotypes.

1. Introduction Rice is the most important food crop in Bangladesh. It is grown

Rice (Oryza sativa L.) is an important cereal crop that feeds more than
50% of the world’s human population [1]. Rice is distinctive in many
countries because of its role in traditional diets and as a major source of
income for a huge number of people. It is grown in over 100 countries,
among which Asia accounts for approximately 87% of global rice
cultivation area and consumes approximately 90% of total rice produced
worldwide [2,3]. Rice is high in energy and accounts for approximately
62% of daily calories and 46% of protein in the average person’s diet [4,
5].
throughout the country and accounts for around 71% of the total
cropped area, which ranks Bangladesh third in the world for rice culti­
vation and overall production after China and India (https://gfmdhaka.
com/news/bangladesh). In Bangladesh, there are three major rice
growing season namely Aus (March–April to July–August), Aman
(June–July to November–December), and Boro (October–November to
April–May), which reflects the vast range of environments and account
for 100% of total rice production [6]. The cultivation area of the three
major rice groups, Aus, Aman, and Boro, was 1.14, 5.87, and 4.75
million hectares, respectively, in fiscal year 2018–19 and produced 2.50,

* Corresponding author. Department of Agronomy, Bangabandhu Sheikh Mujibur Rahman Agricultural University, Gazipur, 1706, Bangladesh.
E-mail addresses: arif@bsmrau.edu.bd, mdarkhan@ttu.edu (Md.A.R. Khan).
1
These authors contributed equally to this work.

https://doi.org/10.1016/j.jafr.2023.100750
Received 8 May 2023; Received in revised form 11 August 2023; Accepted 20 August 2023
Available online 22 August 2023
2666-1543/© 2023 Published by Elsevier B.V. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).
Md.A.R. Khan et al. Journal of Agriculture and Food Research 14 (2023) 100750

13.50, and 18.90 million tons (https://apps.fas.usda.gov/newgainapi/a
pi/report).
Bangladesh is expected to have a total population of 238 million by
2050, with an annual population growth rate of approximately two
million [7,8]. As a result, the country will require extra rice over the next
two decades, but the rice cultivation area in Bangladesh is decreasing
rapidly due to increasing population pressure as well as industrial, res­
idential, and infrastructure development. According to Ref. [9]; the
areas of Aus and Aman experienced a declining tendency, with annual
decreases of − 3.63% and − 0.16%, respectively. Additionally, climate
change has more negative effects on agriculture, especially rice culti­
vation [10]. To adapt to the changing conditions and maximize output,
rice varieties that grow quickly and yield well must be introduced.
As a result, the development of cultivars that grow in a shorter time
and provide higher yields must be prioritized to increase production by
allowing more crops to be grown on the same plot of land each year [11,
12]. Therefore, successful plant breeding efforts entirely depend on
genetic variability and selection expertise. Variability, genetic diversity,
genetic advances, and heritability of traits are the key factors for genetic
improvement [13–15]. The availability of genetic diversity in genotypes
and agronomic traits is crucial for the development of high-yielding and
short-duration varieties because it broadens the gene pool and provides
a chance of selection among the wide range of genotypes to develop a
new desired cultivar [16–18]. The available variability in genotypes can
be assessed using genotypic and phenotypic coefficients of variation,
which are used to partition genetic and environmental variance, ac­
cording to Ref. [19]. The degree of genetic improvement can be char­
acterized by heritability, which reveals the contribution of yield-related
traits to overall production [20]. However, heritability coupled with
genetic advancement will be more helpful in predicting genetic gain
than heritability alone [21]. Principal Component Analysis and biplot
exhibited the level of diversity and degree of association existing be­
tween grain yield and its related component traits [22].
So, selection of promising varieties based on yield components and
genetic variability can increase rice yield and stabilize production under
the worsening situation [23]. Therefore, this study was initiated with
the aim of evaluatoing the 11 rice genotypes considering their genetic
variability of agronomic traits with high yield potential within a shorter
maturity duration in various rice growing seasons of Bangladesh.
2. Materials and method
adopted with three replications containing 11 rice genotypes, including
one more appropriate check cultivar for the particular Aus, Aman, and
Boro seasons (Table 3). The advanced lines used in this experiment were
produced by hybridization procedures from distinct parent lines, and the
variety was obtained from the responsible authority. This parent line
was screened in different experiments on the basis of high yield potential
and short duration. Seeds that had been pre-germinated were planted in
a wet seedbed, and seedlings that were 21, 25, and 35 days old were
transplanted with one seedling per hill, spaced evenly at 25 × 15 cm for
the Aus, Aman, and Boro seasons, respectively, and fertilizer was applied
as [24]. Crop protection measures were taken as required. The necessary
data for determining yield-contributing traits collected during the
maturity stage of the crop are shown in Table 1.
Grain yield. To determine grain yield, 8 square meters of each plot
were harvested, excluding the borders. The grains were then processed,
and the yield was recorded and adjusted to 14% moisture content as
shown in equation (1). Finally, the yield was expressed in metric tons per
hectare.
100 − SMC
Grain yield = × FGW (1)
100 − DMC
Where,
SMC = Sample moisture content, DMC = Desired moisture content,
FGY = Fresh grain weight.
Harvest index (%). Harvest index was estimated by using equation
(2) as follows:
EI
Harvest index (%) = × 100 (2)
BI
The variable “EI” refers to the economic yield, or grain yield, while
“BI” refers to the biological yield, which includes both grain and straw
yield.
Genotypic Variance (σ 2g). Using equation (3) genotypic variance
2
(σ g) was calculated
(MSG − MSE)
σ2 g = (3)
r
Where MSE is the mean square of the error, MSG is the mean square of
genotypes, and r is the number of replications.
Phenotypic Variance (σ 2p). Equation (4) was used to calculate the
phenotypic variance (σ 2p)

The experiment was conducted in the field laboratory of Agronomy σ2 p = σ2 g + σ2 e (4)

at Bangabandhu Sheikh Mujibur Rahman Agricultural University
(BSMRAU), Gazipur-1706 (AEZ-28), during the Aus, Aman, and Boro Where σ 2e is the mean squares of error and σ 2g is genotypic variance.
seasons in 2019–20 at 24◦ 05ˊ North latitude and 90◦ 16ˊ East longitude, Phenotypic and Genotypic Coefficients of Variation (PCV and
and the sea level height is 8.4 m. During the experiment, meteorological GCV). The phenotypic and genotypic coefficients of variation estimates
data such as minimum, maximum, and mean temperature, rainfall, were computed in accordance with [25]; as equations (5) and (6)
sunshine, and humidity were all recorded according to the growing respectively.
season (Table 2). Randomized Complete Block Design (RCBD) was √̅̅̅̅̅̅̅
σ2 p
PCV = × 100 (5)
X
√̅̅̅̅̅̅̅
Table 1 σ2 p
List of studied traits of rice genotypes examined in three seasons with acronym GCV = × 100 (6)
X
and unit of measurement.
The phenotypic variance is represented by σ 2P, while the genotypic
Trait acronym Trait description Unit
variance is represented by σ 2g. The mean of the trait is represented by X.
DM Days to maturity at harvest count The GCV and PCV values were classified as low (0–10%), moderate
PH Plant height at harvest cm
(10–20%), and high (20% and above), following [26].
TT Total tiller hill− 1 at harvest count
ET Effective tiller hill− 1 at harvest count Heritability Estimate. The heritability (broad sense) denoted by
PL Panicle length cm h2B is defined as the ratio of genetic variance (σ 2g) to phenotypic vari­
FG Number of filled grain hill− 1 count ance (σ 2p) according to Ref. [27]; and it can be calculated as equation
UFG Number of unfilled grain hill− 1 Count g (7).
TW 1000-grain weight
GY Grain yield tha− 1 σ2 g
SY Straw yield tha− 1 h2 B = (7)
HI Harvest index percent
σ2 p

2
Md.A.R. Khan et al. Journal of Agriculture and Food Research 14 (2023) 100750

Table 2
Metrological condition during the consecutive three rice season in 2019–20.
Year Month Maximum temperature Average temperature Minimum temperature Precipitation (mm/ Sunshine (h/ Humidity
2019–20 (◦ C) (◦ C) (◦ C) month) month) (%)

Aus March 34 29 20 33.2 279 47

April 37 31 22 293.6 218.5 58
May 34 31 24 542.8 163.5 71
June 32 29 25 857.7 126 87
July 31 29 25 516.9 117 89

Mean 33.6 29.8 23.2 448.84 180.8 70.4

Aman July 31 29 25 516.9 117 89

August 31 30 27 656.3 189 81
September 31 30 27 548.7 255 80
October 31 29 27 833 260 80
November 29 26 23 61.1 336 64

Mean 30.6 28.8 25.8 523.2 231.4 78.8

Boro December 27 25 21 0 372 54

January 27 24 20 3.8 372 51
February 30 26 20 0.2 336 51
March 33 29 24 1 368 62
April 34 31 26 18.8 355 66

Mean 30.2 27 22.2 4.76 360.6 56.8

(https://www.worldweatheronline.com).

√̅̅̅̅̅̅̅
Table 3 GA = K × σ 2 p × h2 B (8)
Rice genotypes for the evaluation of growth and yield attribute in three rice
season. √̅̅̅̅̅̅̅
σ2 p
GAM = K × × h2 B × 100 (9)
Season Genotypic Genotype Genotypic Genotype X
2019–20 code name code name
The intensity of the selection is denoted by the constant K. h2B stands
Aus 1 BU-R-ACC-01 7 BU-R-ACC-07
for heritability, X stands for the mean of the traits, and 2 P is the
2 BU-R-ACC-02 8 BU-R-ACC-08
3 BU-R-ACC-03 9 BU-R-ACC-09 phenotypic standard deviation when K is 5%.
4 BU-R-ACC-04 10 BU-R-ACC-10
5 BU-R-ACC-05 11 BU-R-ACC-11
6 BU-R-ACC-06 12 BINA dhan 19 2.2. Statistical analysis
Aman 1 BU-R-ACC-01 9 BU-R-ACC-09
2 BU-R-ACC-02 10 BU-R-ACC-10 Treatment means were compared using the Least Significant Differ­
3 BU-R-ACC-03 11 BU-R-ACC-11 ence (LSD) test at a significance level of 5% using analytical software
4 BU-R-ACC-04 12 BU dhan 1
statistics 10, according to Ref. [31]. Multivariable analysis was con­
5 BU-R-ACC-05 13 BU dhan 2
6 BU-R-ACC-06 14 BINA dhan 7 ducted using STAR version 2.1.0 [32]. According to [33], replicated data
7 BU-R-ACC-07 15 BRRI dhan 49 were statistically analyzed to determine the sources of variance. The
8 BU-R-ACC-08 16 BRRI dhan 57 cluster plot with heatmap was visualized with the “pheatmap” package;
Boro 1 BU-R-ACC-01 10 BU-R-ACC-10 clustering based on yield and maturity was done through the package
2 BU-R-ACC-02 11 BU-R-ACC-11 “cluster” and the correlation matrix was also created from the package
3 BU-R-ACC-03 12 BRRI dhan 28 “corrplot” in statistical software R (version 3.6.3).
4 BU-R-ACC-04 13 BRRI dhan 29
5 BU-R-ACC-05 14 BRRI dhan 47
6 BU-R-ACC-06 15 BRRI dhan 50 3. Results
7 BU-R-ACC-07 16 BRRI dhan 55
8 BU-R-ACC-08 17 BRRI dhan 67
3.1. Agronomic performances of rice genotypes
9 BU-R-ACC-09 18 BRRI dhan 74

This study characterized the variation in 11 genotypes of rice with

The heritability (broad sense) is determined by the ratio of genetic
variance (σ 2g) to phenotypic variance (σ 2p) [27], and is expressed as a
percentage. The heritability was classified as low (0–30%), moderate
(30–60%), and high (≥60%) based on the categorization proposed by
Ref. [28].
2.1. Measures of genetic advance (GA) and genetic advance as a
percentage of the mean (GAM)
The method of [29] was used to calculate the expected genetic
advance (GA) (equation (8)) and genetic advance as a percentage of the
mean (GAM) (equation (9)) with a selection intensity (K) of 5%. Ac­
cording to Ref. [30]; the genetic advance was then divided into three
categories: low (0–10%), moderate (10–20%), and high (>20%).
suitable cultivars in various growing seasons for morphological and
yield-related traits. A list of traits measured and their ranges are pro­
vided in Table 4. The variation in morphological and yield-related traits
differed significantly among genotypes (P ≤ 0.001) and (P ≤ 0.05),
except for TW in Aman season and HI in Boro season (Table 4). The
findings indicated a potential amount of variability among the geno­
types. The studied genotypes showed the shortest average DM (107.39)
in the Aus season but the lowest PH (105.59 cm) observed in the Boro
season. The higher numbers of TT (16.82), ET (14.11), PL (26.06 cm),
FG (132.56), and UFG (43.31) were recorded in Aus season. The mean
TW (26.60 g) was observed at its maximum in the Aman season, while
the average GY (6.68 tha-1) and HI (60.33%) were recorded at their
highest in the Boro season (Table 4). In the Aus season, the cultivar BINA
dhan 19 matured earlier (96.1 days), and the maximum maturity
duration was found in BU-R-ACC-02 (115.3 days). The lowest PH was

3
Md.A.R. Khan et al. Journal of Agriculture and Food Research 14 (2023) 100750

Table 4
Descriptive statistics and analysis of variance (ANOVA) for eleven evaluated traits related to growth and yield performances of rice genotypes in all three seasons.
Year Traits Minimum Maximum Mean SE SD SS df MS P(>F)
2019–20
a
Aus DM 96.00 116.00 107.39 0.85 5.08 904.56 11 27.36
a
PH 87.00 165.85 119.53 2.72 16.34 9342.80 11 283.00
a
TT 7.00 26.50 16.82 0.82 4.92 846.08 11 24.96
a
ET 4.00 23.00 14.11 0.77 4.64 753.06 11 21.74
a
PL 23.31 28.40 26.06 0.22 1.30 58.70 11 1.69
a
FG 51.90 191.50 132.56 5.40 32.39 36717.99 11 1110.66
a
UFG 16.30 70.50 43.31 2.75 16.49 9514.05 11 282.23
a
TW 20.23 34.00 25.18 0.46 2.76 267.04 11 7.94
a
GY 2.76 7.10 4.87 0.16 0.93 30.48 11 0.92
a
SY 2.43 7.90 4.78 0.19 1.17 47.53 11 1.43
a
HI 35.02 67.29 50.83 1.41 8.48 2516.69 11 75.55
a
Aman DM 106.00 135.00 115.54 6.52 0.94 1995.92 15 44.20
a
PH 100.80 130.20 114.93 7.09 1.02 2359.79 15 50.47
a
TT 7.20 16.60 11.53 2.47 0.36 285.81 15 6.28
a
ET 6.60 16.00 11.00 2.39 0.34 268.36 15 5.91
a
PL 21.40 29.90 24.58 1.68 0.24 132.48 15 2.90
a
FG 81.90 182.30 128.11 23.94 3.46 26947.86 15 597.38
b
UFG 7.00 39.90 16.46 6.83 0.99 2195.45 15 46.92
TW 3.12 32.80 26.60 5.37 0.78 1357.01 15 29.04 NS
a
GY 4.21 7.29 5.65 0.56 0.08 14.56 15 0.31
b
SY 2.40 7.24 4.65 0.81 0.12 31.19 15 0.60
b
HI 24.89 62.91 50.19 9.73 1.40 4452.47 15 91.99
a
Boro DM 128.00 150.00 140.63 0.80 5.87 1824.59 17 35.48
a
PH 80.28 116.28 105.59 1.26 9.26 4542.05 17 88.70
a
TT 8.60 20.60 13.61 0.31 2.28 275.11 17 5.37
a
ET 7.20 18.20 12.07 0.28 2.04 219.76 17 4.29
a
PL 21.04 26.79 23.71 0.17 1.27 85.50 17 1.68
a
FG 79.80 164.30 118.29 2.84 20.89 23135.37 17 451.56
a
UFG 3.10 53.50 22.18 1.68 12.31 8032.89 17 155.76
a
TW 20.15 30.45 26.14 0.41 3.05 492.84 17 9.64
a
GY 3.27 9.76 6.68 0.17 1.24 81.13 17 1.59
a
SY 3.89 9.27 5.86 0.14 1.06 59.86 17 1.17
HI 51.55 65.79 60.33 0.39 2.87 435.77 17 8.31 NS

Abbreviations: SE, standard error; SD, standard deviation; SS, sum of square; MS, mean sum of square; Pr, probability value, DM, days to maturity; PH, plant height; TT,
total tillers hill-1; ET, effective tillers hill-1; PL, panicle length; FG, filled grains panicle-1; UEG, unfilled grains panicle-1; TW, test weight; GY, grain yield; SY, straw
yield; HI, harvest Index.
** significant at 0.01 level.
a
Significant at 0.001 level.
b
Significant at 0.05 level.

observed in BU-R-ACC-10 (108.1 cm), and the maximum TT (24.7) and
ET (25.5) were found in the genotype BU-R-ACC-04. The trait PL was
highest in BU-R-ACC-06 (27.6 cm), followed by BU-R-ACC-09 (27.5 cm)
and BU-R-ACC-10 (27.4 cm). The highest FG was found in BU-R-ACC-11
(174.2), and the lowest UFG was in BINA dhan 19 (22) followed by BU-
R-ACC-09 (26.1). The genotype BU-R-ACC-09 showed the highest TW
(27.9 g), followed by BU-R-ACC-01 (27.8 g), while the highest GY was
produced by the genotype BU-R-ACC-08 (6.2 tha-1), which represented
the statistical similarity with the BU-R-ACC-11 (5.8 tha-1), BU-R-ACC-01
(5.7 tha-1), BU-R-ACC-04 (5.6 tha-1), BU-R-ACC-05 (5.2 tha-1), BU-R-
ACC-07 (5.2 tha-1), BU-R-ACC-09 (4.9 tha-1) and BU-R-ACC-10 (4.8
tha-1). The genotype BU-R-ACC-02 showed the highest SY (7.6 tha-1),

Table 5
Performance of genotypes at harvest as affected by growth and yield component traits genotypes in Aus season.
Genotypes DM PH TT ET PL FG UFG TW GY SY HI
ef b a a d g bc a ab ab
BU-R-ACC-01 105.0 126.1 24.7 20.5 25.9- 95.4f 35.2 27.8 5.7 6.8 45.7c-e
BU-R-ACC-02 115.3a 127.6b 17.8b-e 15.8b-d 25.3de 112.9d-g 39.7bc 26.3a-c 4.3b-d 7.6a 36.2e
BU-R-ACC-03 113.3b 116.0de 9.7g 5.7g 25.6cd 136.4b-e 62.1a 23.4c-e 3.0d 4.6cd 39.7de
BU-R-ACC-04 112.3b 161.7a 19.0b-d 16.5a-c 26.2b-d 129.0d-f 33.1bc 25.1a-d 5.6ab 5.6bc 49.6a-d
BU-R-ACC-05 104.3f 112.6ef 21.3ab 18.3ab 25.9cd 85.5g 37.4bc 25.43a-c 5.2ab 4.4cd 54.2a-c
BU-R-ACC-06 106.0e 116.8c-e 14.2d-g 11.2ef 27.6a 130.1c-f 48.8ab 27.5ab 4.4bc 2.9e 59.8a
BU-R-ACC-07 105.0ef 116.3de 14.3d-g 11.7d-f 23.9e 164.6a-c 62.1a 24.5b-d 5.2ab 4.1de 55.7a-c
BU-R-ACC-08 104.0f 119.6cd 20.8a-c 17.7a-c 26.9abc 169.7ab 41.9a-c 22.1de 6.2a 4.8cd 56.5ab
BU-R-ACC-09 108.7d 122.7bc 13.3e-g 11.0f 27.5ab 146.8a-d 26.1c 27.9a 4.9ab 4.0de 55.1a-c
BU-R-ACC-10 110.3c 108.1f 18.8b-d 15.7b-e 27.4ab 140.1a-e 60.4a 20.9e 4.8ab 5.0cd 49.5b-d
BU-R-ACC-11 108.33d 117.1c-e 16.2c-f 13.7c-f 25.3de 174.2a 50.9ab 23.7de 5.8a 4.7cd 55.7a-c
BINA dhan 19 96.1g 89.4g 11.6fg 11.7d-f 25.2de 106.0e-g 22.0c 27.5ab 3.2cd 2.9e 52.3a-c

Lsd (0.05) 1.1 5.9 4.9 4.6 1.4 34.8 20.5 3.2 1.3 1.2 10.3
CV% 0.6 2.9 17.5 19.4 3.1 15.5 27.6 7.6 16.5 15.1 12.1

Data in column with the same letter are not significantly different at the 0.05 level by least significant difference test.
Abbreviation: DM, days to maturity; PH, plant height; TT, total tillers hill− 1; ET, effective tillers hill− 1; PL, panicle length; FG, filled grains panicle− 1; UEG, unfilled
grains panicle− 1; TW, test weight; GY, grain yield; SY, straw yield HI, harvest Index; LSD, least significant difference; CV, Coefficient of variation.

4
Md.A.R. Khan et al. Journal of Agriculture and Food Research 14 (2023) 100750

followed by BU-R-ACC-01 (6.8 tha-1), while the genotype BU-R-ACC-06
exhibited a higher HI (59.8%) (Table 5). In the case of Aman season, the
BU-R-ACC-11 showed the shortest DM (107 days), which was statisti­
cally similar with the BU-R-ACC-05 (109 days). The lowest PH was
found in BU-R-ACC-10 (105.4 cm), followed by BU-R-ACC-03 (107.4
cm). BU dhan 2 exhibited higher TT (15.6) and ET (14.6) followed by
BINA dhan 7 (14.4), and (13.9). The higher PL was observed in BU dhan
2 (27.4) and FG in BU-R-ACC-11 (158.8). The lowest UFG was observed
in BU-R-ACC-10 (9.3), and there was no significant difference in the
value of TW in Aman season. The cultivar BRRI dhan 49 produced the
highest GY (6.9 tha-1), which was statistically similar with the BU dhan
2, BU-R-ACC-05, and BU-R-ACC-06 (6.1 tha-1) (Table 6). The BRRI dhan
49 showed the highest SY (6.2 tha-1) while the HI was higher in BU-R-
ACC-01 and BU-R-ACC-10 (57.5%) followed by BU-R-ACC-05 (57.1%)
and BU-R-ACC-11(56.8%) (Table 6). The lowest DM during the irrigated
Boro season was recorded in BRRI dhan 28 (129.3 days), which was
statistically comparable with BRRI dhan 74 (131.3 days). The lowest PH
was found in BRRI dhan 29 (97.7 cm), followed by BU-R-ACC-05 (105.3
cm) and BRRI dhan 55 (105.7 cm). The genotypes BRRI dhan 50
exhibited higher TT (18.3) and ET (15.8). The highest PL was found in
BRRI dhan 47 (25.8 cm), while a higher number of FG was observed in
BU-R-ACC-11 (152.6). The lowest number of UFG was found in BRRI
dhan 67 (5.9), followed by (8.9), while BU-R-ACC-09 showed the
highest TW (30.2 g) among the genotypes. The highest grain yield was
found in BU-R-ACC-07 (8.8 tha-1), and there were no statistical differ­
ences among the genotypes: BU-R-ACC-06 (8.4 tha-1), BU-R-ACC-02 (7.9
tha-1), BU-R-ACC-01 (7.9 tha-1), BU-R-ACC-04 (7.7 tha-1), BU-R-ACC-08,
and BU-R-ACC-09 (7.7 tha-1). The highest SY and HI were found in BRRI
dhan 29 (8.5 tha-1) and BU-R-ACC-05 (63.4%), respectively (Table 7).
3.2. Estimation of genetic parameters
The genotypic and phenotypic variances (δ2g and δ2p), genotypic and
phenotypic coefficients of variation (GCV and PCV), broad sense heri­
tability (h2B) and genetic advance as a percentage of the mean (GAM)
for yield and yield related agronomic traits as shown in Table 8. The trait
FG displayed the widest range of genotypic and phenotypic variations
among the traits followed by the UFG, PH, and DM, while GY displayed
the least variation in all rice growing seasons. It was found that the PCV
was higher than the GCV in all of the characters, indicating that the
environment has the effects on the expression of characters [34]. There
was a high to medium estimation of GCV and PCV were obtained for
most of the traits across all the growing seasons. However, the lowest
GCV values were found for DM (4.83, 5.59 and 4.11), PL (3.90, 5.13 and
4.39) and HI (12.17, 9.37 and 1.94) in Aus, Aman and Boro season,
respectively. In case of PCV, the traits DM (4.87, 5.75 and 4.24) and PL
(4.99, 6.92 and 5.46) showed the lowest values in all growing seasons.
There was some variation in terms of h2B during the three seasons and
most of the traits exhibited high to moderate heritability except for HI
(24.05 and 16.44) in Aman and Boro season which is defined as lower.
Most of the studied traits showed moderate to high genetic advances
(GA) and genetic advances as a percentage of mean (GAM) for all
growing seasons while the traits FG recorded the highest (GA) 42.52,
22.56 and 26.53 respectively in all the three seasons, but the lowest for
GY in Aus (1.08) and Aman (0.46) whereas the HI was lowest in Boro
season (0.98). The GAM was highest for SY (49.28) in Aus, ET (28.25) in
Aman and UFG (84.70) in Boro season while the lowest value was for PL
(6.27) in Aus, TW (4.79) in Aman and HI (1.62) for Boro season
respectively.
3.3. Principal component analysis
We were able to determine the characters that were responsible for
genotypic variation within the group using principal component anal­
ysis (PCA). PCA was carried out using 11 traits for various genotypes in
different growing seasons, whereas eigenvalues less than one should not
be taken into consideration for any factors of PCA. In this study, the first
four principal components (PC) had eigen values greater than one in Aus
and Boro seasons, but it was found in the first five principals in Aman
season and their cumulative contribution to the variation from three
season assessment was 93.76%, 89.3%, and 89.52%. The variability
percent of the PC1 was the highest at 31.3%, 30.9% and 40.52% in Aus,
Aman and Boro seasons, respectively. The positive and negative loadings
indicate the existence of a positive and negative connection between the
variable and the component [35]. PC1 was contributed positively by the
three characters such as TT (0.482), ET (0.464) and SY (0.445) and PC2
contributed mostly by two characters like DM (0.502) and UFG (0.473)
in the genotypes of Aus season. In case of Aman season, the PC1 posi­
tively contributed by three characters namely TT (0.434), ET (0.435)
and UFG (0.0393) while PC2 contributed by the TT (0.318), TW (0.331)
and HI (0.367). Among the rice genotypes of Boro season, the PC1
positively contributed by TT (0.395) and ET (0.388) while PC2 was
availed of by TW (0.408) only. On the other hand, the traits namely FG
(− 0.128), UFG (− 0.104) and HI (− 0.158) contributed negatively to the

Table 6
Performance of genotypes at harvest as affected by growth and yield component traits genotypes in Aman season.
Genotypes DM PH TT ET PL FG UFG TW GY SY HI

BU-R-ACC-01 110.3fg 114.5b-d 14.7ab 14.6a 23.5cd 96.6f 15.6b-e 29.1a 5.7b-e 4.2b-d 57.5a
BU-R-ACC-02 117.0de 116.7bc 9.4e-g 9.1c-e 23.9c 95.6f 21.2a-c 28.4a 4.8e 5.3ab 47.6a-d
BU-R-ACC-03 110.7fg 107.4de 8.3g 7.9de 23.5cd 140.0a-d 10.5de 25.6a 5.1c-e 4.8bc 51.9a-c
BU-R-ACC-04 121.3b 126.3a 11.1c-e 10.4b-d 23.3cd 129.7a-e 13.8b-e 29.6a 5.8bc 5.3ab 52.1a-c
BU-R-ACC-05 109.0gh 114.4b-d 13.3a-c 12.8ab 24.4c 115.1c-f 10.1e 29.9a 6.1ab 4.6bc 57.1a
BU-R-ACC-06 111.3fg 112.3c-e 12.1b-d 11.07bc 26.7 ab 154.7ab 17.1b-e 29.1a 6.1b 4.9a-c 55.4ab
BU-R-ACC-07 112.7f 115.4bc 10.9c-f 10.67bc 24.5c 139.4a-d 20.1a-d 29.2a 5.5b-e 4.4b-d 55.8ab
BU-R-ACC-08 112.3f 112.8cd 12.3b-d 11.3bc 26.7 ab 138.6a-d 22.7ab 24.3a 5.2b-e 5.2ab 51.0a-d
BU-R-ACC-09 116.0e 110.1c-e 8.3fg 7.6e 25.2bc 127.8b-e 12.9c-e 21.3a 4.9de 4.4b-d 52.8a-c
BU-R-ACC-10 110.7fg 105.4e 10.8c-g 10.1c-e 24.4c 144.4a-c 9.3e 24.3a 5.9bc 4.4b-d 57.5a
BU-R-ACC-11 107.0h 112.73c-e 10.3d-g 9.93c-e 24.2c 158.8a 12.3c-e 26.3a 5.4b-e 4.1b-d 56.8a
BU dhan 1 116.7de 125.7a 11.1c-e 11.0bc 25.1bc 113.3d-f 18.0b-e 25.6a 5.9bc 4.9a-c 43.8a-d
BU dhan 2 121.3b 120.8ab 15.6a 14.6a 27.4a 133.0a-e 27.7a 29.9a 6.1ab 5.0a-c 42.3b-d
BINA dhan 7 118.7cd 107.4de 14.4ab 13.9a 24.3c 103.6ef 17.4b-e 25.7a 5.7b-d 3.8cd 41.4cd
BRRI dhan 49 133.0a 115.9bc 10.1d-g 9.9c-e 21.7d 141.6a-d 17.5b-e 22.9a 6.9a 6.2a 42.1b-d
BRRI dhan 57 120.7bc 120.8ab 11.6c-e 11.2bc 24.5c 117.3c-f 17.2b-e 24.6a 5.7b-d 3.1d 37.7d

Lsd (0.05) 2.6 7.3 2.6 2.5 1.9 30.3 9.6 9.5 0.9 1.4 13.9
CV% 1.4 3.8 13.6 13.6 4.6 14.2 35.1 21.3 9.4 17.5 12.0

Data in column with the same letter are not significantly different at the 0.05 level by least significant difference test.
Abbreviation: DM, days to maturity; PH, plant height; TT, total tillers hill− 1; ET, effective tillers hill− 1; PL, panicle length; FG, filled grains panicle− 1; UEG, unfilled
grains panicle− 1; TW, test weight; GY, grain yield; SY, straw yield; HI, harvest index; LSD, least significant difference; CV, Coefficient of variation.

5
Md.A.R. Khan et al. Journal of Agriculture and Food Research 14 (2023) 100750

Table 7
Performance of genotypes at harvest as affected by growth and yield component traits genotypes in Boro season.
Genotypes DM PH TT ET PL FG UFG TW GY SY HI
cd a-d b-d bcd ab d-h a-c a-c fg
BU-R-ACC-01 143.7 107.7 14.7 13.1 25.1 90.2h 16.2 28.9 7.9 5.0 60.9a-e
BU-R-ACC-02 149.0a 108.2a-d 12.7c-e 11.3cde 23.3d-g 112.9c-g 16.6d-h 28.6b-d 7.9a-c 5.8ef 57.7de
BU-R-ACC-03 149.7a 111.7a-c 10.4e 8.6f 23.8b-f 141.3ab 25.1cd 29.2ab 7.6bc 4.8g 61.2a-e
BU-R-ACC-04 141.0ef 108.8a-d 14.3b-d 12.3b-e 22.4gh 107.6d-h 10.3f-h 27.5c-e 7.7a-c 4.8g 61.7a-d
BU-R-ACC-05 137.0gh 105.3b-e 15.8ab 14.3ab 24.0b-e 93.4gh 14.8d-h 28.8a-c 7.4bc 4.2g 63.4a
BU-R-ACC-06 145.7bc 109.6a-d 11.8de 10.0ef 24.7a-c 124.3b-e 29.7c 27.2de 8.4ab 5.1fg 62.1a-c
BU-R-ACC-07 144.7cd 115.5a 12.5c-e 11.5c-e 22.6fg 133.7a-c 40.8ab 24.8fg 8.8a 5.1fg 63.2ab
BU-R-ACC-08 142.3de 110.6a-c 13.0b-e 11.7b-e 24.9ab 133.3a-c 17.4d-g 24.5gh 7.7a-c 5.1fg 59.8a-e
BU-R-ACC-09 147.7ab 114.0ab 12.3de 11.5c-e 24.5b-d 127.9b-d 12.5e-h 30.2a 7.7a-c 4.9fg 60.8a-e
BU-R-ACC-10 137.3gh 101.4de 14.4b-d 13.2bc 23.1e-g 98.1f-h 20.5c-f 23.1h 7.0c 4.9fg 58.9b-e
BU-R-ACC-11 139.0fg 105.2c-e 12.0de 10.5d-f 23.4d-g 152.6a 21.8c-e 26.5e 7.5bc 4.8g 60.7a-e
BRRI dhan 28 129.3i 110.1a-c 15.3a-c 13.2bc 22.5gh 115.7c-f 22.4c-e 23.2h 4.8d 6.9b-d 59.2a-e
BRRI dhan 29 144.7cd 97.7e 14.6b-d 13.3a-c 24.5a-d 124.3b-e 30.6bc 21.1i 5.3d 8.5a 61.4a-e
BRRI dhan 47 135.0h 107.1a-d 13.1b-e 11.5c-e 25.8a 126.7b-d 30.1bc 26.1ef 5.3d 7.5ab 58.7c-e
BRRI dhan 50 140.7ef 82.9f 18.3a 15.8a 22.9e-g 102.6e-h 50.4a 20.2i 4.8d 6.6c-e 57.6de
BRRI dhan 55 138.0g 105.7b-e 12.4c-e 11.2c-e 23.6c-g 111.0d-h 25.2cd 27.5c-e 5.3d 7.6ab 59.2a-e
BRRI dhan 67 135.3h 110.5a-c 14.0b-d 12.1b-e 24.4b-d 139.0ab 5.9h 23.5gh 4.6d 7.5bc 62.3a-c
BRRI dhan 74 131.3i 88.5f 13.4b-d 12.1b-e 21.3h 94.5f-h 8.9gh 29.5ab 4.7d 6.2de 57.1e

Lsd (0.05) 2.4 8.7 2.9 2.6 1.3 22.1 10.7 1.6 1.2 0.9 4.4
CV% 1.1 4.9 13.1 12.9 3.3 11.3 29.2 3.6 10.4 9.9 4.4

Data in column with the same letter are not significantly different at the 0.05 level by least significant difference test.
Abbreviation: DM, days to maturity; PH, plant height; TT, total tillers hill− 1; ET, effective tillers hill− 1; PL, panicle length; FG, filled grains panicle− 1; UEG, unfilled
grains panicle− 1; TW, test weight; GY, grain yield; SY, straw yield; HI, harvest index; LSD, least significant difference; CV, Coefficient of variation.

Table 8
Estimation of genetic variances and attributes for plant growth and yield-related traits studied among the rice genotypes for different growing season.
Year Traits Variance components Coefficient of variations h2B(%) Genetic advance
2019–20
δ2g δ2p GCV (%) PCV (%) GA GAM (%)

Aus DM 26.95 27.36 4.83 4.87 98.51 10.61 9.88

PH 270.63 283.00 13.76 14.07 95.63 33.14 27.73
TT 16.35 24.96 24.04 29.70 65.50 6.74 40.08
ET 14.24 21.74 26.75 33.05 65.51 6.29 44.59
PL 1.03 1.69 3.90 4.99 61.05 1.63 6.27
FG 687.90 1110.67 19.79 25.14 61.94 42.52 32.08
UFG 139.19 282.22 27.24 38.79 49.32 17.07 39.41
TW 4.29 7.92 8.22 11.18 54.09 3.14 12.46
GY 0.50 0.92 18.23 24.61 54.89 1.08 27.83
SY 1.09 1.43 27.35 31.26 76.54 1.89 49.28
HI 38.29 75.56 12.17 17.10 50.67 9.07 17.85

Aman DM 41.72 44.20 5.59 5.75 94.40 12.93 11.19

PH 31.16 50.47 4.86 6.18 61.74 9.04 7.86
TT 3.81 6.28 16.94 21.75 60.65 3.13 27.17
ET 3.67 5.91 17.40 22.08 62.09 3.11 28.25
PL 1.59 2.90 5.13 6.92 54.97 1.93 7.84
FG 267.63 597.38 12.77 19.08 44.80 22.56 17.61
UFG 13.60 46.92 22.39 41.60 28.98 4.09 24.83
TW 3.15 25.89 6.67 19.12 12.16 1.27 4.79
GY 0.12 0.31 7.81 12.24 40.72 0.46 10.26
SY 0.18 0.60 11.50 20.89 30.31 0.48 13.04
HI 22.12 91.99 9.37 19.11 24.05 4.75 9.47

Boro DM 33.33 35.48 4.11 4.24 93.94 11.53 8.20

PH 61.44 88.70 7.42 8.92 69.27 13.44 12.73
TT 2.19 5.37 10.86 17.02 40.72 1.94 14.28
ET 1.88 4.29 11.36 17.17 43.79 1.87 15.49
PL 1.08 1.68 4.39 5.46 64.59 1.72 7.26
FG 273.71 451.56 13.99 17.96 60.61 26.53 22.43
UFG 113.80 155.76 48.10 56.28 73.06 18.78 84.70
TW 8.76 9.64 11.32 11.88 90.86 5.81 22.23
GY 1.28 1.59 21.15 23.57 80.51 2.09 39.08
SY 0.95 1.17 20.82 23.07 81.48 1.82 38.71
HI 1.37 8.31 1.94 4.78 16.44 0.98 1.62

Abbreviation: DM, days to maturity; PH, plant height; TT, total tillers hill− 1; ET, effective tillers hill− 1; PL, panicle length; FG, filled grains panicle− 1; UEG, unfilled
grains panicle− 1; TW, test weight; GY, grain yield; SY, straw yield; HI, harvest index; σ 2g, genotypic variance; σ 2p, phenotypic variance; PCV, phenotypic coefficient of
variation; GCV, genotypic coefficient of variation; h2B, heritability percentage, GA, genetic advance; GAM, genetic advance as percentage of the mean.

variation in PC1 in very negligible way, whereas the traits TT (− 0.167), (− 0.212) and HI (− 0.307) contributed negatively to PC1 and DM
ET (− 0.246), PL (− 0.120) and TW (− 0.384) contributed negatively to (− 0.532), PH (− 0.191), GY (− 0.191) and SY (− 0.325) to PC2 respec­
PC2 in the genotypes of Aus season. In case of Aman season, the traits FG tively. Most traits namely DM (− 0.297), PH (− 0.0383), FG (− 0.251),

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Md.A.R. Khan et al. Journal of Agriculture and Food Research 14 (2023) 100750

TW (− 0.290), GY (− 0.372) and HI (− 0.252) negatively impacted vari­
ability in PC1 and all traits, except TW contributed negatively in PC2
among the rice genotypes in Boro season (Table 9).
3.4. Trait-specific genotypic relationship through biplot
Biplot analysis provides a visual comparison among genotypes and
an association among traits based on various traits by measuring the
angle of trait vector from the origin. The more acute angle (<90◦ )
Table 9
Eigen values, proportion of the total variance represented by first four principal
components, cumulative per cent variance and component loading of different
characters in rice for yield and yield component traits.
Season Parameters PC1 PC2 PC3 PC4 PC5
(2019-20
between the vector of traits exhibited a stronger positive relation and the
more obtuse angle (>90◦ ) correlated negatively with the trait relation
The longer the vector from the origin, the more the contribution of the
parameter to the divergence and wider variability was caused by the
genotypes that were the furthest away from the origin of trait vectors
[36]. The traits PL, FG HI and TW contribute less variability to the first
two PCs while the traits PL, TW, DM, and also PH contribute less vari­
ability to PC1 and PC3 and the GY was more acutely angled with the
traits TT, ET, and PL for the first three PCs in the Aus season (Fig. 1-b). In
the Aman season, the traits PL, FG, and SY showed less variability in PC1
and PC2 while the trait PL contributed less variability in PC1 and PC3.
The GY showed more acute angle with PH in PC1 and PC2 and with TW
in PC1 and PC3 (1c-d). The traits PL, FG, and UFG contributed less
variability in the first three PCs while the traits GY showed an acute
angle with the traits PL, DM, HI, and PH in the first three PCs for the
genotypes of the Boro season (1e-f).

Aus EigenValues 3.44 2.53 2.25 1.2 0.88 3.5. Correlation among yield and related traits
Variability 31.3 23.09 20.44 10.9 8.03
(%)
The relationship between various plant characteristics related to
Cumulative 31.3 54.39 74.83 85.73 93.76
(%) yield and its constituent parts is measured by correlation coefficient
DM 0.195 0.502 − 0.136 − 0.35 0.073 analysis, which can be used to guide selection for genetic improvement.
PH 0.354 0.171 − 0.004 − 0.467 − 0.393 The linear, perfect positive linear, and perfect negative linear relation­
TT 0.482 − 0.167 0.134 0.188 0.19 ships are represented by the r-values of 0, +1, and − 1, respectively. Low,
ET 0.464 − 0.246 0.106 0.217 0.124
PL 0.047 − 0.12 0.182 − 0.625 0.672
moderate, and strong positive linear correlations are shown by values
FG − 0.128 0.298 0.475 − 0.105 − 0.312 ranging from 0 to 0.3, 0.3 to 0.7, and 0.7 to 1, respectively, while low,
UFG − 0.104 0.473 0.224 0.26 0.178 moderate, and strong negative linear correlations are shown by values
TW − 0.033 − 0.384 − 0.365 − 0.276 − 0.354 ranging from 0 to − 0.3, − 0.3 to − 0.7, and − 0.7 to − 1 [13]. In the
GY 0.368 − 0.08 0.444 0.004 − 0.252
present study, correlation analysis indicated a significant positive as­
SY 0.445 0.217 − 0.236 0.12 − 0.032
HI − 0.158 − 0.319 0.507 − 0.131 − 0.143 sociation between different agronomic traits. There was a strong posi­
tive correlation between ET and TT (r = 0.97**) while a medium
Aman EigenValues 3.40 2.47 1.46 1.36 1.13
Variability 30.9 22.46 13.24 12.4 10.29
positive relationship exists between PH with DM (r = 0.6**); GY with TT
(%) (r = 0.68**) as well as ET (r = 0.66**); and also SY with DM (r =
Cumulative 30.9 53.37 66.6 79 89.3 0.53**). The traits SY showed moderate positive correlation with PH (r
(%) = 0.47**), TT (r = 0.41*) and ET (r = 0.36*); GY with PH (r = 0.41*)
DM 0.258 − 0.532 0.04 − 0.048 0.09
and HI (r = 0.52**) in Aus season (Fig. 2a). On the contrary, there was a
PH 0.323 − 0.191 − 0.086 0.059 − 0.467
TT 0.434 0.318 − 0.019 − 0.149 0.221 moderate to strong negative association between UFG and ET (r =
ET 0.435 0.304 0.011 − 0.226 0.178 − 0.37*); TW and FG (r = − 0.50**); UFG (r = − 0.47**); HI and DM (r =
PL 0.154 0.307 − 0.044 0.632 0.211 − 0.47**), SY (r = − 0.73**) (Fig. 2a). In the Aman season, there exist
FG − 0.212 − 0.043 − 0.537 0.273 0.416 strong positive significant correlations between ET and TT (r = 0.98**)
UFG 0.393 − 0.061 0.04 0.505 − 0.055
TW 0.241 0.331 − 0.327 − 0.1 − 0.475
while a lower to moderately significant association was observed in PH
GY 0.27 − 0.191 − 0.409 − 0.381 0.392 with DM (r = 0.37**); PL with TT (r = 0.30*); UFG with PH (r = 0.40**);
SY 0.058 − 0.325 − 0.526 0.139 − 0.249 PL (0.32*); GY with DM (r = 0.38**); PH (r = 0.32*); TT (r = 0.35*); ET
HI − 0.307 0.367 − 0.383 − 0.121 − 0.177 (r = 0.36*); and SY with PH (r = 0.35*); GY (r = 0.43**). The analysis
Boro EigenValues 4.45 1.86 1.56 1.13 0.83 revealed moderately negative significant correlations between FG and
Variability 40.52 16.95 14.14 10.34 7.56 TT (r = − 0.33*), ET (r = − 0.38**); HI and DM (r = − 0.56**), UFG (r =
(%) − 0.42**) (Fig. 2b). In the Boro season, there was a strong positive and
Cumulative 40.52 57.47 71.61 81.96 89.52
(%)
significant relationship between TT and ET (r = 0.97**) and moderate to
DM − 0.297 − 0.065 − 0.376 − 0.322 0.355 lower significant correlations were found between FG with PH (r =
PH − 0.383 − 0.089 0.019 0.341 − 0.147 0.46**), PL (r = 0.36**); TW with PH (r = 0.31*); GY with DM (r =
TT 0.395 0.145 − 0.326 0.243 0.008 0.61**), PH (r = 0.53**), TW (r = 0.43**); and finally HI with PH (r =
ET 0.388 0.167 − 0.334 0.244 0.038
0.33*), GY (r = 0.41**) (Fig. 2c). Furthermore, moderate to lower
PL − 0.164 − 0.322 − 0.146 0.408 0.696
FG − 0.251 − 0.515 0.091 − 0.048 − 0.341 negative significant correlations were observed among the traits such as
UFG 0.125 − 0.355 − 0.454 − 0.453 − 0.042 GY with SY (r = − 0.65***), TT (r = − 0.33*) and ET (r = − 0.29*); PH
TW − 0.29 0.408 0.287 − 0.047 0.296 with TT (r = − 0.31*), ET (r = − 0.32*); DM with SY (r = − 0.34*), TT (r
GY − 0.372 0.231 − 0.372 − 0.124 − 0.028 = − 0.29*) and ET (r = − 0.29*) and also TW with SY (r = − 0.43**), UFG
SY 0.258 − 0.471 0.274 0.098 0.205
HI − 0.253 − 0.052 − 0.331 0.509 − 0.345
(r = − 0.46***), TT (r = − 044***) and ET (r = − 0.42**) in the boro rice
growing season (Fig. 2c).
Abbreviation: PC, principal component; DM, days to maturity; PH, plant height;
TT, total tillers hill− 1; ET, effective tillers hill− 1; PL, panicle length; FG, filled
4. Discussion
grains panicle− 1; UEG, unfilled grains panicle− 1; TW, test weight; GY, grain
yield; SY, straw yield; HI, harvest index. Five principal components were
extracted for eleven yield growth related agronomic traits with different Eigen The study was actually focused on the variations of yield and yield
value for different rice growing season. The first four components had Eigen related agronomic traits and their contribution to yield for the evalua­
value greater than one Aus and Boro season whereas first five components tion of rice genotypes. In the present study, the analysis of variance
exhibited Eigen value more than one in Aman season. The value of loading indicated considerable variation on observed characters and there is
indicated the contribution of different traits to the variability. possibility of genetic improvement through selection and hybridization

7
Md.A.R. Khan et al.
for this significant yield and its attributing traits on tested genotypes. As
can be seen from this study, some genotypes mature earlier than others.
Early maturing indicates a short life cycle and is considered a positive
character for rice improvement. Early maturing varieties are advanta­
geous in rice based cropping system because they grow faster during the
vegetative phase and thus more competitive with weeds, evacuating the
land early for the next crop, escaping insect pest attack, and utilizing less
water [37] & [38]. Some varieties namely BU-R-ACC-05, BU-R-ACC-01,
BU-R-ACC-10 and BU-R-ACC-11 matured earlier in our experiment for
three seasons and were highly desirable. This finding complied with the
result of [39] who found a significant difference in days to maturity
among 24 tested rice genotypes. Plant height generally depends on their
number and length of internodes. It is indirectly related to yield by
attributing effective assimilate partitioning at low expense and being
advantageous in protection against lodging [40]. Therefore, short to
moderate plant height is desirable when breeding for high-yielding va­
rieties [41]. In this experiment, it was found that all of the high-yielding
genotypes were of moderate height for all growing seasons. This in­
dicates that moderate plant height is desirable when breeding for
8
Journal of Agriculture and Food Research 14 (2023) 100750
Fig. 1. PCA biplot view of the genotype with the
contribution of trait by the length of vector. Fig. 1
(a–b), Fig. 1(c–d), Fig. 1(e–f) indicated the Aus, Aman
and Boro season’s biplot view respectively. The biplot
view showed the interrelationships among traits. The
first three PCs exhibited maximum variability each
and cumulatively to the total variation. The angle
between the vectors of two traits approximates the
correlation coefficients between the traits. The name
of genotypes is presented in Table 1. Abbreviation:
DM, days to maturity; PH, plant height; TT, total til­
lers hill− 1; ET, effective tillers hill− 1; PL, panicle
length; FG, filled grains panicle− 1; UEG, unfilled
grains panicle− 1; TW, test weight; GY, grain yield; SY,
straw yield; HI, harvest Index.
high-yielding varieties. Our results comply with the findings of [37].
Tillering ability in rice is closely related to the panicles produced per
unit area and finally impacts on the varietal yield [42]. Therefore, ge­
notypes with higher productive tillering ability could be chosen by
breeders for the improved high yielding variety. The aforementioned
higher number of productive tillers that depends on number of tillers
was observed in the proposed genotypes such as BU-R-ACC-01 and
BU-R-ACC-05 in Aus; BU-R-ACC-01 and BU dhan 2 in Aman;
BU-R-ACC-05 and BRRI dhan 50 in Boro season. Similar results were
observed by Ref. [43] who stated that tiller number hill− 1 varied among
the varieties where PR 121 produced maximum effective tillers (11.7
hill− 1) and least effective tillers (9.3 hill− 1) have been recorded in PAU
201. They also expressed that the number of effective tillers was
responsible for yield enhancement (r = 0.56) in the rice. Similarly [44],
strongly implied that the quantity and quality of tillers determine the
yield of rice. The panicle length is a key factor that directly influences
the yield, and the variation in panicle length might be due to the vari­
ation in genetic makeup of different rice genotypes [45]. In the present
study, the genotypes BU-R-ACC-06, BU-R-ACC-08, BU-R-ACC-09, and
Md.A.R. Khan et al.
Fig. 2. Pearson’s correlation analysis results of various genotypes among the eleven yield and growth related traits.
Abbreviation: DM, days to maturity; PH, plant height; TT, total tillers hill− 1; ET, effective tillers hill− 1; PL, panicle length; FG, filled grains panicle− 1; UEG, unfilled
grains panicle− 1; TW, test weight; GY, grain yield; SY, straw yield; HI, harvest Index.
BU-R-ACC-10 exhibited the highest panicle length in all three seasons
along with BU-R-ACC-01 in Boro season. Credence to this research
finding has been laid by Ref. [46] who showed that panicle length varied
with the planting technique and that the length of the rice panicle
significantly influenced the grain yield. The filled grain panicle− 1 is one
of the important yield attributing traits. An increase in the number of
filled grains could be attributed to efficient translocation of carbohy­
drates from the sources to the spikelets (sinks) which consequently leads
to an increase in grain yield [47]. The genotypes BU-R-ACC-11
contributed a higher number of filled grains panicle− 1 in all three sea­
sons [38]. observed variation in fertile grain number per panicle in
various rice experiments in Khumaltar, Lalitpur of Nepal. Thousand
grain weight influences grain yield greatly, which is determined by both
grain size and grain filling rate in rice [48]. The result of the present
study showed that 1000 grain weight varied significantly among the
tested genotypes. Similarly [49], reported that thousand grain weight
was mainly governed by genetic factors and that significant differences
were found among twenty rice genotypes in Nepal. The traits filled with
grain number panicle− 1, 1000 grain weight, plant height, panicle length,
grain panicle− 1, seed setting rate, lengthy growth period, low panicle
number per unit area, and low seed length/width ratio have been found
to be responsible for high grain production [50]. The result of our study
showed that the genotypes BU-R-ACC-08 (4.9 tha-1) and BU-R-ACC-11
(4.7 tha-1) produced the highest grain yield in Aus season,
9
Journal of Agriculture and Food Research 14 (2023) 100750
BU-R-ACC-05 (4.8 tha-1) and BU-R-ACC-06 (4.8 tha-1) in Aman season
and BU-R-ACC-07 (7 tha-1) and BU-R-ACC-06 (6.7 tha-1) in Boro season.
These findings are similar to those of [51,52] who found differences in
grain yield traits among the 12 rice genotypes and reported that the
genetic makeup of the crop is the most likely cause of variation in grain
yields. The harvest index is a vital characteristic of physiological sig­
nificance that reflects the transfer of a dry matter to the economic parts
[53]. In the present study, the BU-R-ACC-01, BU-R-ACC-04,
BU-R-ACC-05, BU-R-ACC-06, BU-R-ACC-08, BU-R-ACC-09 and
BU-R-ACC-11 genotypes pose the maximum harvest index in rice
growing seasons. Similarly [54], found significant variation in harvest
index among the seven promising lines of rice and also reported that
higher yields can be obtained through a higher rate of partitioning of
photosynthates from source to sink. The traits assessed in rice accessions
showed significant variations in the variance components among the
studied genotypes that indicate the existence of considerable genetic
variability among themselves with regard to the 11 agronomic traits.
Results of the present study closely agree with the earlier reports of [55]
who found significant variations among 40 rice accessions for 12
quantitative traits. The magnitude of the differences between the
genotypic and phenotypic coefficients of variation indicates the envi­
ronmental influence on any character [56]. We found that PCV was
slightly to moderately higher than the GCV for all the traits and high to
moderate GCV and PCV values were observed for the most important
Md.A.R. Khan et al.
yield related traits in all the growing seasons except DM, PL and HI [57].
also found the higher GCV and PCV values for most important
yield-associated characteristics such as the number of grains panicle− 1,
panicles hill− 1, and tillers hill− 1. Higher heritability as well as high
genetic advance give better clues than individual parameters which
pointed out non-additive gene action and suggested that selection would
be effective [58]. In this study, most of the yield attributing traits
showed moderate to high heritability (h2B) where the traits PH (95.63
and 33.14), FG (61.94 and 42.52), DM (98.51 and 10.61) and UFG
(49.32 and 17.07) exhibited high to moderate h2B with high to moderate
GA in Aus season. In case of Aman season, moderate to high h2B and GA
were exhibited by the traits DM (94.40 and 12.93) and FG (44.80 and
22.56) and in Boro, the traits DM (93.94 and 11.53), FG (60.61 and
26.53) and UFG (73.06 &18.78) showed moderate to high h2B and GA.
This finding complied with those of [58] who found high heritability
combined with moderate to high GA for the traits of number of tillers per
meter square, days to flowering, and grain yield. The principal compo­
nent analysis has been widely used for various purposes, and it em­
phasizes the significance of larger contributors to the overall variation at
each axis of differentiation [22]. In the present study, principal
component analysis showed a high level of variability and diversity in
the studied rice genotypes for different seasons, considering eleven
agronomic traits. Five principal components were extracted, with ei­
genvalues ranging from 0.88 to 3.44. The first three principal compo­
nents explained 74.83%, 66.6% and 71.61% of the total variation with
respect to the Aus, Aman and Boro seasons. Generally, traits included in
the first principal components, especially traits with high loading scores,
are important for separating the genotypes [16]. The total variability in
the genotypes is due to cumulative effect of a number of traits rather
than the small contribution of each character. In the first principal
component, traits like TT, ET, and SY contributed more to the total
variation (31.30%) due to their high loading. The traits DM and UFG
played a significant role in the second major component, which was
responsible for 23.09% of variation in aus season. In the case of Aman
season, the traits TT, ET and UFG were the greatest contributors to the
variation (30.90%) in the first principal component, whereas the traits
TT, TW and HI were important in the second principal component,
which contributed 22.46% of the total variation. The traits TT and ET
contributed greater total variation (40.52%) in the first principal
component due to their high loading values while the trait TW was
important in the second principal component, which contributed
16.95% of the total variation in Boro season. Similarly [22], explained
64% of the total variability using the first three principal components
among 64 rice genotypes based on 7 morphological traits. Similar results
were found in the work of [59] where 86% of the total variation was
explained by the first five principal components respectively.
The biplots were an excellent tool for visualizing genotype-by-trait
and revealing the interrelationships among traits on the basis of multi­
ple traits and identifying genotypes or groups of genotypes that can be
candidates for introduction as best genotypes [60]. The PCA biplot
revealed that the genotypes BINA dhan 19, BU-R-ACC-03, BU-R-ACC-01,
BU-R-ACC-02, BU-R-ACC-06 and traits DM, TT, ET, UFG and BY were
dispersed and away from the origin and had high genetic variability
between the PC1 and PC2 in Boro season. In the aman season, the BRRI
dhan 49 and BU dhan 2 cultivars and the traits HI, TT, ET, DM are placed
dispersedly in between the PC1 and PC2 and have more genetic vari­
ability. Another in Boro season, the genotypes BRRI dhan 29, BRRI dhan
50, BRRI dhan 74, BRRI dhan 47, BU-R-ACC-03 and the traits like TW,
GY, PH, FG, SY, TT and ET away from the origin between the PC1 and
PC2 indicate high variability. The traits GY showed strong positive re­
lations with the traits of TT, ET, and PL in Aus season; with the traits of
PH and TW in Aman season and the traits of PL, DM, HI, and PH in Boro
season for PC1, PC2 and PC3 which are measured through the acute
angle between these traits. The genotypes BU-R-ACC-01, BU-R-ACC-05
in Aus season, BU dhan 1, BRRI dhan 57 in Aman season and
BU-R-ACC-01, BU-R-ACC-02 and BU-R-ACC-05 contributed as the best
10
Journal of Agriculture and Food Research 14 (2023) 100750
genotypes under the studied characters. The results are closely related to
the findings of [61] who stated that genotypes and traits closer to the
origin and closer to each other indicate a low contribution to the
divergence. Correlation coefficient analysis measures the mutual rela­
tionship and also the degree and direction of relationships between
various traits including grain yield [62,63]. Degree of correlation be­
tween the traits is important in plant breeding as it can be used as a tool
for indirect selection. A positive correlation between desirable charac­
ters is favorable because it helps in simultaneous improvement of both
characters [64]. In the present study, the correlation analysis revealed
significant positive and negative correlations among the studied traits,
especially with grain yield. The grin yield was positively and correlated
with the plant height, total tillers, effective tillers and harvest index in
Aus season; days to maturity, plant height, total tillers, effective tillers
and straw yield in Aman season; plant height, harvest index, test weight
and days to maturity in Boro season. Similar results were reported by
Ref. [65] who stated that yield was positively correlated with number of
panicle plant− 1, number of grains panicle− 1 and fertility percentage
[54]. also observed a similar high positive significant correlation be­
tween grain yield, filled grains per panicle with spikelet fertility, harvest
index and flag leaf area.
5. Conclusion
The development of rice genotypes rests on genetic aspects of its
quantitative traits, which are essential in the development of new va­
rieties. The overall results indicated that all the genotypes showed
variations in yield and yield-related traits. The traits plant height, total
tillers, effective tillers, filled grains, unfilled grains, and test weight
showed moderate to high heritability and genetic advance in percentage
of the mean. The rice genotypes, namely, BU-R-ACC-08 and BU-R-ACC-
11, produced better yields in Aus season, BU-R-ACC-05 and BU-R-ACC-
06 in Aman season, and BU-R-ACC-07 and BU-R-ACC-06 in Boro season,
as well as the BU-R-ACC-05, BU-R-ACC-01, BU-R-ACC-10, and BU-R-
ACC-11 genotypes mature earlier in all three season. Therefore, these
high-yielding genotypes could be selected to obtain a better yield for the
particular rice growing season. Further studies are recommended to
execute research focusing on metabolic and molecular profiling for a
better understanding of the mechanisms.
Authors’ contributions
Md. Arifur Rahman Khan (MARK), Apple Mahmud (AM), Md.
Nahidul Islam (MNI), Uttam Kumar Ghosh (UKG), Md. Saddam Hossain
(MSH), Conceptualization: MARK; Investigation: MARK, AM; Data
Analysis and interpretation: MARK, AM, MNI; Writing—original draft
preparation: MARK, AM, MNI; Writing—review and editing: MARK,
AM, MNI, UKG, MSH; Supervision and Project administration: MARK;
Funding acquisition: MARK.
Declaration of competing interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influence
the work reported in this paper.
Data availability
Data will be made available on request.
Acknowledgements
The authors thank the Ministry of Science and Technology, People’s
Republic of Bangladesh, for financial support regarding this study.
Md.A.R. Khan et al. Journal of Agriculture and Food Research 14 (2023) 100750

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