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The study was conducted using thirty six upland rice genotypes in 2017 at Gojeb and Guraferda;
Southwestern Ethiopia to classify and identify groups of similar genotypes and thereby estimate the
genetic difference between clusters of the genotypes, the experiment was laid down in 6 × 6 simple
lattice design. The combined analysis of variance over the two locations revealed that the genotypes
showed highly significant (P≤0.01) differences for all the characters studied, except for days to 50%
heading, panicle weight, thousand seed weight, lodging incidences, leaf blast and brown spot.
Similarly genotype × location interactions revealed highly significant (P≤0.01) differences for panicle
shattering and grain yield and significant (P≤0.05) differences for days to 85% maturity, plant height,
number of fertile tillers per plant, number of unfilled spikelets per panicle and biomass yield. The
squared distance (D2) analysis grouped the 36 genotypes in to four clusters. This makes the
genotypes moderately divergent. The Chi-square (x2) test showed that all inter-cluster squared
distances was highly significant. The principal component analysis revealed that four principal
components have accounted for 70.54% of the total variation. The present study revealed that
number of panicles per meter square and harvest index can be considered for selection.
Key-words: upland rice, genetic diversity, cluster analysis, principal component analysis
INTRODUCTION
Rice belongs to the genus Oryza within the grass family Rice is also an important staple food crop in many African
Gramineae (Poaceae). There are about 25 species of countries. It is mostly cultivated in West Africa (Smith,
Oryza, of these, only two species are cultivated, namely 2001). It has been the most rapidly growing food source
Oryza sativa and Oryza glaberrima. Rice (Oryza sativa L.) across the continent. However, local production is mainly
is the most important food crop and energy source for insufficient to meet consumption needs. Annual rice
about half of the world’s population (Manjappa et al., production in Sub-Saharan Africa (SSA) is estimated to be
2014). More than 3.5 billion people in the world depend on 14.5 million metric tons, smallholder farmers produce
rice for more than 20% of their daily calories (Khare et al., most of this rice, in contrast, Africa‘s rice consumption is
2014). Rice (Oryza sativa L.) is grown in more than 117 about 21 million metric tons creating a deficit of about 6.5
countries across all habitable continents covering a total million metric tons per year valued at US$ 1.7 billion that
area of about 163 million hectares with a global production is imported annually, overall, imported rice accounts for
of about 740 million metric tons ( FAOSTAT, 2014). Asia roughly 31 percent of Sub-Saharan Africa’s local rice
is the leader in rice production accounting for about 90% consumption (AATF, 2013).
of the world's production. Over 75% of the world supply is
consumed by people in Asian countries and thus, rice is of *Corresponding Author: Awel Beshir, Southern
immense importance to the food security of Asia (IGC, Agricultural Research Institute, Bonga Agricultural
2014). Research Center,
E-mail: awelbeshir234@gmail.com Tel: +251916357008
Investigating the Genetic Diversity in Upland Rice (Oryza Sativa L.) Genotypes Evaluated at Gojeb and Guraferda, Southwest Ethiopia
Awel et al. 866
Cultivation of rice in Ethiopia is a recent phenomenon it breeders in choosing the right type of parents for
was started first at Fogera and Gambella Plains in the early hybridization program. The divergence can be studied by
1970’s (Tamirat et al., 2017) and grown under widely using technique D2 statistics developed by Mahalanobis
varying conditions of altitude and climate. It is an important (1936). Estimation of genetic diversity is also an important
cereal crop cultivated in different parts of the country next factor in knowing the source of genes for a particular trait
to teff, maize, wheat and sorghum (MoA, 2010). within the available genotypes. Appropriate selection of
Considering the importance and potential of the crop, it has the parents is essential to be used in crossing nurseries to
been recognized by the government as “the new enhance the genetic recombination for potential yield
millennium crop of Ethiopia” to attain food security. The increase (Islam, 2004). the principal component analysis
potential area for rice production in Ethiopia is estimated helps researchers to distinguish the significant relationship
to be about 30 million hectares (5 million hectares highly between traits. This is a multivariate analysis method that
suitable and about 25 million hectares suitable) (Ministry aims to explain the correlation between a large set of
of Agriculture, 2010). The average rice productivity in variables in terms of a small number of underlying
Ethiopia is about 2.8 t ha-1 (Central Statistical Authority, independent factors. The main objective of this study is to
2017), which is much lower than that of the world’s estimate the magnitude of genetic divergence present in
average (4.4 t ha-1) (FAOSTAT, 2016). This low the 36 upland rice genotypes by using cluster analysis and
productivity of rice in Ethiopia is attributed to many factors cluster analysis-PCA-based methods for the selection of
such as shortage of genetic improved varieties, lack of parents in hybridization programme.
recommended crop management practices, lack of pre-
and post-harvest management technologies and lack of
awareness on its utilization (Tesfaye et al., 2005). MATERIALS AND METHODS
One of the most important approaches to rice breeding is Description of the Experimental Areas: The experiment
hybridization. Parents’ choice is the first step in plant was conducted during the 2017 main rainy season in two
breeding programs through hybridization (Joshi et al., locations namely, Gojeb and Guraferda, Southwestern
2004). Genetic diversity is the most important tool for plant Ethiopia.
Experimental Materials: In this experiment, 33 upland rice genotypes, obtained from two different sets of variety trials
conducted by rice breeding section of Fogera National Rice Research and Training Center (FNRRTC) and three released
varieties (NERICA-12, NERICA-4 and Adet), a total of 36 upland rice genotypes were used (Table 2).
Investigating the Genetic Diversity in Upland Rice (Oryza Sativa L.) Genotypes Evaluated at Gojeb and Guraferda, Southwest Ethiopia
ART16-9-1-9-2-1-1-B-1-1 2016/17 PVT Africa Rice ART15-19-5-4-1-1-1-B-1-1 2016/17 NVT Africa Rice
Center Center
ART16-9-4-18-4-2-1-B-1-1 2016/17 PVT Africa Rice ART16-5-9-22-2-1-1-B-1-2 2016/17 NVT Africa Rice
Center Center
ART16-9-4-18-4-2-1-B-1-2 2016/17 PVT Africa Rice ART16-21-4-7-2-2-2-B-2-2 2016/17 NVT Africa Rice
Center Center
ART16-9-6-18-1-1-2-B-1-1 2016/17 PVT Africa Rice ART16-9-16-21-1-2-1-B-1- 2016/17 NVT Africa Rice
Center 1 Center
ART16-9-9-25-2-1-1-B-2-1 2016/17 PVT Africa Rice ART15-13-2-2-2-1-1-B-1-2 2016/17 NVT Africa Rice
Center Center
ART16-9-9-25-2-1-1-B-2-2 2016/17 PVT Africa Rice ART15-16-45-1-B-1-1-B-1- 2016/17 NVT Africa Rice
Center 2 Center
ART16-9-29-16-1-1-1-B-1- 2016/17 PVT Africa Rice ART16-5-10-2-3-B-1-B-1-1 2016/17 NVT Africa Rice
1 Center Center
ART16 15-10-1-1-B-1-B-B- 2016/17 PVT Africa Rice ART16-4-1-21-2-B-2-B-1-2 2016/17 NVT Africa Rice
1 Center Center
ART16 15-10-1-1-B-1-B-B- 2016/17 PVT Africa Rice NERICA-12 Released Africa Rice
2 Center Center
ART16-13-11-1-2-B-2-B-2- 2016/17 PVT Africa Rice NERICA-4 Released Africa Rice
1 Center Center
ART16-16-1-14-3-1-1-B-1- 2016/17 PVT Africa Rice Adet Released Africa Rice
2 Center Center
PVT = Preliminary Variety Trial and NVT = National Variety Trial, FNRRTC = Fogera National Rice Research and Training
Center.
Investigating the Genetic Diversity in Upland Rice (Oryza Sativa L.) Genotypes Evaluated at Gojeb and Guraferda, Southwest Ethiopia
Awel et al. 868
Genetic divergence analysis Xp = the subject’s score on observed variable p.
The principal component analysis was computed using the The mean squares due to genotype × location interactions
correlation matrix of SAS version 9.3 (SAS, 2014) to have differed significantly for panicle shattering, grain yield
examine the relationships among the quantitative (kg ha-1), days to 85% maturity, plant height, fertile tillers
characters that are correlated with each other by per plant, unfilled spikelets per panicle and biomass yield.
converting into uncorrelated characters called principal This significant difference of genotype × location
components. Below is the general formula to compute interactions indicates the differential response of
scores on the first component extracted (created) in a genotypes to the two locations for these characters.
principal component analysis. Ogunbayo et al. (2014) found significant genotype x
location interaction effects for days to maturity, plant height
PC1 = b11(X1) + b12 + ⋯ b1p (Xp) and grain yield. Tefera et al. (2017) also reported
significant genotype x location interaction effects for
Where: PC1 = the subject’s score on principal component biomass yield, days to maturity, fertile tillers per plant, plant
1 (the first component extracted); height, panicle length and grain yield.
b1p = the regression coefficient (or weight) for observed
variable p, as used in creating principal
component1
Table 3: Mean squares of combined analysis of variance for 18 characters of 36 upland rice genotypes
evaluated in 2017 main cropping season across two locations
Characters MSL(1) MSG(35) MSGxL(35) MSE(60) CV
DH 15.34ns 9.55ns 7.01ns 6.10 3.04
DM 108.51** 22.61** 13.91* 8.29 2.46
PH 190.44** 79.67** 19.72* 11.75 4.11
PL 0.75ns 2.98** 1.51ns 1.08 4.82
TTPP 26.52** 6.14** 2.67ns 2.61 13.21
FTPP 18.20** 6.11** 3.67* 2.09 14.97
FSPP 123.58ns 324.73** 165.40ns 155.29 12.73
USPP 27.04** 5.74** 3.96* 2.33 13.78
PW 15.80** 2.56ns 1.82ns 1.76 8.78
Pan/m2 30.25ns 120.29** 47.55ns 44.95 14.36
BY 4033402.78** 1128445.63** 686545.63* 413372.20 15.68
HI 0.0001ns 0.0067** 0.0036ns 0.0027 14.64
Investigating the Genetic Diversity in Upland Rice (Oryza Sativa L.) Genotypes Evaluated at Gojeb and Guraferda, Southwest Ethiopia
TSW 55.95** 10.48ns 7.06ns 6.68 9.06
+LB 11.61 ns 26.55 ns 25.02ns 16.78 19.13
+BS 72.35* 24.90ns 21.97ns 16.27 15.08
$LI 0.001ns 0.50ns 0.42ns 0.32 20.10
$PSht 0.92ns 2.47** 2.12** 0.29 22.71
GY 1898539.52** 288979.76** 226956.50** 111151.56 12.68
The numbers in the brackets indicates degree of freedom, + = mean squares are based on arc sin, $ = mean squares are
based on square root transformation * = significant at (P≤0.05), ** = significant at (P≤ 0.01), MSL = mean squares of
locations, MSG = mean squares of genotypes, MSGxL = mean square of genotype x location interaction, MSE = mean
squares of error, CV = coefficient of variation. DH = days to 50% heading, DM = days to 85% maturity, PH = plant height,
PL = panicle length, TTPP = total tillers per plant, FTPP = fertile tillers per plant, FSPP = filled spikelets per panicle, USPP
= unfilled spikelets per panicle, PW = panicle weight, Pan/m2 = panicles per meter square, BY = biomass yield, HI =
harvest index, TSW = thousand seed weight, LB = leaf blast, BS = brown spot, LI = lodging, PSht = panicle shattering,
GY = grain yield (kg ha-1).
Multivariate Analysis
Cluster analysis
The genotypes were partitioned into four distinct groups relationship among themselves as compared to the other
based on their similarities in characteristics (Table 4 and clusters. Therefore, it could be expected that genotypes
Figure 1); this makes the genotypes to be moderately within a cluster are less genetically different with each
divergent. Cluster I contained the highest number of other, but are diverse from the genotypes belonging to
genotypes 17 (47%), followed by clusters II that contained other clusters. Hossain et al. (2015) grouped thirty three
15 (42%) genotypes. It also comprising two checks droughts tolerant rice (Oryza sativa L.) genotypes into
(NERICA-12 and Adet). These genotypes may be seven clusters. Cluster I and II were comprised of the
regarded as having the overall characteristics of these maximum number of genotypes (8) in each followed by
checks. In contrast, cluster III and IV had the smallest cluster V containing 5 genotypes. Khare et al. (2014)
number of genotypes, consisted of 3 (8%) and 1 (3%), clustered sixty upland rice accessions into seven groups.
respectively. This cluster analysis showed that the rice Cluster III contained the highest 14 accessions, followed
genotypes were originated from different sources. by clusters I comprised 11.
Genotypes falling in a particular cluster indicate their close
Investigating the Genetic Diversity in Upland Rice (Oryza Sativa L.) Genotypes Evaluated at Gojeb and Guraferda, Southwest Ethiopia
Awel et al. 870
Figure 1: Dendrogram indicating the genetic relationship of 36 upland rice genotypes evaluated over the two
locations at Southwestern Ethiopia.
44.05
Similarity
62.70
81.35
100.00
- 1 -1 - 2 - 2 - 2 - 1 -2 - 1 - 1 - 2 - 1 - 2 -2 - 1 - 2 - 2 - 1 -1 - 1 - 1 - 2 - 1 -2 - 1 12 - 1 - 1 - 1 - 2 - 1 - 1 et - 1 -4 - 2 - 2
B - B B- 1 B- 1B- B B -1 B- 2 B- 1 B- 1B- B B - 1B -1 B- 1 B- 1 B- 1B- B B -1 B- 1B- B B- B B- 1B -1 B- 2 B- 2 B- 1CA B - 1B -1 B- 1 B- 1 B- 2 B- 1 A d B- 1I C A B- 1 B- 2
1- 1- 1- 2- 1- 1- 1- 2- 1- 1- 2- 1- 1- 1- 1- 1- 1- 2- 1- 1- 2- 1- 1- 1- I 2- 1- 1- 1- 2- 1- 1- R 1- 2-
1- 2- 1- 2- 2- 1- 1- B - B- 1- 1- 1- 2- 2- B - B- B- B- B- 1- B- 1- 1- B - ER 1- 1- 1- B - B - 1- 2-1- NE - 1-2- 2-
-4- - 4- - 2- - 2- - 4- - 2- - 2- - 2- - 1- -1- - 1- - 3- - 2- - 1- - 1- - 1- -3- -1- - 2- -1- - 2- - 3- - 2- - 1- N -1- - 1- - 2- - 3- - 2- -2- - -B -
36 18 - 2 46 18 25 22 21 - 1 - 2 18 14 16 21 - 1 25 - 2 1 0 - 4 21 12 25 18 9-129- 162- 330- 2 1- 113- 1 - 1- 9 5- 1- 4- 7
10- - 4- 3- 217- - 4- - 9- - 9- - 1- - 10 - 33 - 6- - 1- 14- 16- - 10 11- - 1016- 229-19- 5- 1- - 5- - 9- - 7- 2 2 2 - 1 -1 - - 9 - 4 1
8- 6- 9 5- 1 0- 6- 9 6-9 6- 5 6-4 15 6- 9 6- 9 - 16 - 9- -9- 15 6- 6- 5 9- 9- 5- 1 6- 4 - 21 6- 9 - 17 - - 4
- 9 - 9 9- 1 6-5 - 13 6- T16 5- 1616-
2
T15RT1RT1 15 1R T1RT 1RT1 T 1T16R T1RT 1T16T 16T 16T 16 16- 1 T1T16T16R T1 T1 T16RT1T16 T16T1616- T 1T16R T1 R T1 RT
T R R R T R A
A R A AA R A A A A A R A A A R A R A R A RA R T A A R A R A A A R A AR A R A RA R A A R A AR A
Ge notype s
Comparison of genotype performance among clusters the lowest cluster mean value for plant height, panicle
length, number of fertile tillers per plant, number of filled
The mean value of the 12 characters for each cluster group spikelets per panicle, biomass yield, harvest index and
is presented in Table 5. Cluster I was characterized by the grain yield (kg ha-1). Cluster IV produced the highest
highest cluster mean estimate for plant height, number of cluster mean values for days to 85% maturity, panicle
filled spikelets per panicle, number of unfilled spikelets per length, number of total tillers per plant, number of fertile
panicle and harvest index. However, it produced the tillers per plant, number of panicles per meter square,
lowest cluster mean estimate for the number of total tillers biomass yield and grain yield (kg ha-1). However, it
per plant. Cluster II was characterized by having higher produced the lowest cluster mean values for the number
cluster mean values for harvest index and panicle of unfilled spikelets per panicle and panicle shattering.
shattering. However, it produced the lowest cluster mean Therefore, these typical characteristics in each clusters
estimate for days to 85% maturity and number of panicles may be used for the variety development program through
per meter square. Cluster III was characterized by having selection or/and hybridization.
Investigating the Genetic Diversity in Upland Rice (Oryza Sativa L.) Genotypes Evaluated at Gojeb and Guraferda, Southwest Ethiopia
Int. J. Plant Breed. Crop Sci. 871
Genetic divergence analysis from crosses involving parents selected from cluster three
and four followed by cluster two and four. The minimum
The standardized Mahalanobis D2 statistics showed that squared distance was found between cluster two and three
there is a high genetic distance and highly significant (D2 = 25.41) followed by cluster one and two (D2 = 29.40)
variation at P<0.01 and P<0.05 among the four clusters and cluster one and four (D2 = 83.21), indicating that
(Table 6). The maximum squared distance was found genotypes in these clusters were not genetically diverse or
between cluster three and four (D2 = 313.86) followed by there was little genetic diversity between these clusters.
cluster two and four (D2 = 189.27). Maximum genetic This signifies that crossing of genotypes from these
recombination is expected from the parents selected from clusters might not give higher heterotic value in F1 and a
divergent clusters groups. Therefore, maximum narrow range of variability in the segregating F 2 population.
recombination and segregation of progenies is expected
Table 6: Intra and inter-cluster values of generalized square distance (D 2) among four clusters constructed from
36 upland rice genotypes
Clusters Cluster I Cluster II Cluster III Cluster IV
Cluster I 1.50ns 29.40** 97.29** 83.21**
Cluster II 1.75ns 25.41** 189.27**
Cluster III 4.96ns 313.86**
Cluster IV 0.00ns
2
X = 24.72 and 19.67 at 1% and 5% probability level, respectively, ** = highly significant, ns = non-significant, bold values
are intra-cluster distance.
Principal component analysis principal component (PC2) contributed 14.94% of the total
variation. The major contributing characters for the
The principal component analysis revealed four principal variation in the second principal components (PC2) were
components PC1, PC2, PC3 and PC4 with eigenvalues days to 85% maturity, plant height, number of total tillers
greater than one (Table 7). They have accounted for per plant, number of fertile tillers per plant, number of filled
70.54% of the total variation among genotypes for the spikelets per panicle, number of unfilled spikelets per
twelve quantitative characters. Khare et al. (2014) reported panicle, biomass yield and panicle shattering.
that the combination of the first four principal components
accounted for 77.13% of the total variation of all the In the same way, 11.62% of the total variability among the
characters. The relative magnitude of eigenvectors from tested genotypes accounted for the third principal
the first principal component (PC1) was 35% showing that component (PC3) originated from variation in plant height,
all characters except plant height and number of filled panicle length and number of total tillers per plant. The
spikelets per panicle had high loading and most fourth principal component (PC4) contributed 8.98% of the
contributing characters for the total variation. The second total variation. The number of unfilled spikelets per panicle
Investigating the Genetic Diversity in Upland Rice (Oryza Sativa L.) Genotypes Evaluated at Gojeb and Guraferda, Southwest Ethiopia
Awel et al. 872
and harvest index expressed the highest loads in principal Therefore, the above mentioned characters with high
component four (PC4). The positive and negative weight positive or negative loads contributed more to the variation
shows the presence of positive and negative correlation and they were the ones that most differentiated the
trends between the components and the variables. clusters.
Table 7: Eigenvectors and Eigenvalues of the first four principal components (PCs) for 12 characters of 36 upland
rice genotypes
Characters PCA 1 PCA 2 PCA 3 PCA 4
DM 0.47 0.40 0.11 0.09
PH -0.11 0.54 0.70 0.01
PL 0.59 0.12 0.63 -0.13
TTPP 0.43 -0.55 0.55 -0.02
FTPP 0.79 -0.41 0.21 -0.08
FSPP 0.19 0.71 -0.17 -0.01
USPP -0.65 0.35 0.25 -0.48
Pan/m2 0.80 -0.12 -0.06 0.14
BY 0.50 0.45 -0.03 -0.001
HI 0.69 -0.11 -0.21 -0.35
PSht -0.71 -0.29 0.27 -0.02
GY 0.89 0.24 -0.13 0.12
Eigen value 4.55 1.94 1.51 1.17
Proportion (%) 35 14.94 11.62 8.98
Cumulative (%) 35 49.94 61.56 70.54
DM = days to 85% maturity, PH = plant height, PL = panicle length, TTPP = number of total tillers per plant, FTPP =
number of fertile tillers per plant, FSPP = number of filled spikelets per panicle, USPP = number of unfilled spikelets per
panicle, Pan/m2 = number of panicles per meter square, BY = biomass yield, HI = harvest index, PSht = panicle shattering,
GY = grain yield (kg ha-1).
The combined analysis of variance revealed that, the There is no conflict of interest among the authors or
genotypes were significantly different for all the characters anybody else.
studied, except days to 50% heading, panicle weight,
thousand seed weight, lodging incidence and reaction to Acknowledgements
major rice diseases (leaf blast and brown spot). This
indicates the existence of a considerable amount of First I would like to thank Southern Agricultural
variation among the tested genotypes. The genotype × Research Institute for financing the study and Jimma
location interaction effects were also significant for days to University College of Agriculture and Veterinary
85% maturity, plant height, number of fertile tillers per Medicine for hosting the study. I gratefully acknowledge
plant, number of unfilled spikelets per panicle, biomass the immense contribution made by Bonga Agricultural
yield, panicle shattering and grain yield, indicates that Research Center during the execution of the research
differential response of genotypes under the two locations work. It provided me necessary facilities that have made
for these characters. The genotypes were partitioned into my work smooth. I am also highly indebted to Fogera
four distinct groups based on their similarities in Rice Research and Training Center for the provision of
characteristics; this makes the genotypes to be moderately necessary materials.
divergent. There were statistically approved differences
between clusters. The maximum squared distance was
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Investigating the Genetic Diversity in Upland Rice (Oryza Sativa L.) Genotypes Evaluated at Gojeb and Guraferda, Southwest Ethiopia