GENETIC VARIABILITY AND CHARACTER ASSOCIATION IN

BREAD WHEAT (Triticum aestivum L.) GENOTYPES AT JAMMA

AND GEREGERA, ETHIOPIA

M. Sc. THESIS

AHMED GETACHEW

FEBRUARY, 2017

JIMMA, ETHIOPIA
GENETIC VARIABILITY AND CHARACTER ASSOCIATION IN
BREAD WHEAT (Triticum aestivum L.) GENOTYPES AT JAMMA
AND GEREGERA, ETHIOPIA

M.Sc. Thesis

Submitted to the School of Graduate Studies, College of Agriculture

and Veterinary Medicine, Jimma University in Partial Fulfillment of
the Requirements for the Degree of Master of Science in Plant
Breeding

Ahmed Getachew

February, 2017

Jimma, Ethiopia
 Jimma University College of Agriculture and Veterinary Medicine
Thesis Submission Request Form (F-05)
Name: Ahmed Getachew ID No. RM1265/2007

Program of Study: Plant Breeding

Tittle: Genetic Variability and Character Association in Bread Wheat (Triticum aestivum L.)
Genotypes at Jamma and Geregera, Ethiopia

I have completed my thesis research work as per the approved proposal and it has been evaluated
and accepted by my advisers. Hence I hereby kindly request the Department to allow me to
present the findings of my work and submit the thesis.

Ahmed Getachew
Name and signature of student
We, the thesis advisers have evaluated the contents of this thesis and found to be satisfactory,
executed according to the approved proposal, written according to the standards and format of
the University and is ready to be submitted. Hence, we recommend the thesis to be submitted.

Dr. Sentayehu Alamerew __________________ ___________________

Major advisor Signature Date

Dr. Fisseha Worede ___________________

Co-advisor Signature Date

Internal Examiner (If depends on the Verdict)

Name Signature Date
Decision/ Suggestion of Department Graduate council (DGC)

Chairperson, DGC Signature Date

___________________ ___________________ ___________________
Chairperson, CGS Signature Date
___________________ ___________________ ___________________
 DEDICATION

I dedicate this thesis manuscript to my beloved mother Belay Adinew, my father Getachew
Assen, my son Abubekr Ahmed and to my uncle Seid Hassen.

I
 STATEMENT OF THE AUTHOR

First of all, I declare that this thesis is my original work and all sources of materials used for this
thesis have been duly acknowledged. This thesis has been submitted in partial fulfillment of the
requirements for M.Sc. degree in plant breeding at Jimma University and is deposited at the
University Library to be available to borrowers under the rules of the library. I solemnly declare
that this thesis is not submitted to any other institution anywhere else for the award of any
academic degree, diploma, or certificate.

Brief quotations from this thesis are allowable without special permission, provided that accurate
acknowledgement of source is made. Requests for permission for extended quotation from or
reproduction of this manuscript in whole or part of it may be granted by the head of the major
department or the Dean of School of Graduate Studies, when in his or her judgment the proposed
use of the material is in the interest of scholarship. In all other instances, however, permission
must be obtained from the author.

Name: Ahmed Getachew Assen Signature _____________________

Place: Jimma University, Jimma
Date of submission: ________________

II
 BIOGRAPHICAL SKETCH

The author, Ahmed Getachew Assen, was born on April 19, 1988 in Wogdie Woreda, South
Wollo Zone, Amhara Regional State, Ethiopia. He attended his primary education in Demasiko
Elementary School from 1995 to 2001.He attended his Junior and secondary school education at
Fita, from 2002 to 2003. He attended his Secondary School from 2004 to 2005 in Wogdie high
School and he completed his Preparatory School in Borena preparatory School from 2006 to
2007 G.C.

In 2008, he joined at Jimma University College of Agriculture and Veterinary Medicine and
graduated with Bachelor of Science degree in Crop science in 2010. He then joined in Amhara
region, Borena Woreda Agricultural Office in 2011, as an expert in Irrigation and Vegetative -
Fruit development department work division till he joined the School of graduate studies of
Jimma University College of Agriculture and Veterinary Medicine for Master of Science in
Agriculture majoring in Plant Breeding in November 2014.

III
 ACKNOWLEDGEMENTS

First of all, I would like to thank Almighty Allah for anything, and all praise is due to Allah, the
Eternal, the Creator, the One and the worship.

I would like to express my deepest gratitude to my major advisor Dr. Sentayehu Alamerew, and
my co-advisor Dr. Fisseha Worede for their guidance, support, continued follow up, and
encouragement throughout the entire periods of the research project and thesis work and I
appreciated them for valuable input in this paper for their unreserved technical comments and
suggestions.

I would be highly indebted to Amhara Regional Agricultural Bureau and Borena Woreda Office
of Agriculture for giving me the chance to pursue my higher education. I would also like to thank
Jimma University College of Agriculture and Veterinary Medicine, for creating a learning
environment and providing resident and computer facilities during my stay in the campus. My
thanks extend to Sirinka Agricultural Research Center for providing me experimental area and
materials.

I am deeply indebted to my father, Getachew Assen and my mother, Belay Adinew, My sister
Sofya Ali, my uncle, Seid Hassen, my brother, Omer Getachew, Mr. Demssew Abebe and Mr.
Moges Adinew for their encouragement and financial support. Special thanks are reserved for
my sister Hayat Tadye for providing me Laptop. Special thanks are reserved for Mr. Tilahun
Abera head of Borena Woreda Rural development and Agricultural Office.

I would like to thank Mr. Zerihun Tadesse, researcher at Kulumsa Agricultural Research Center.
I also appreciate the support provided of Mr. Nuru Ahmed, Mr. Eyeberu and Mr. Mekonen field
assistants at Sirinka Agricultural Research Center.

IV
 LIST of ACRONYMS and ABBREVIATIONS

ANOVA Analysis of Variance

bp base pair

CSA Central Statistical Agency

CV Coefficient of Variation

FAO Food and Agricultural Organization of the United Nations

GAIN Global Agricultural Information Network

GCV Genotypic Coefficient of Variation

LSD Least Significance Difference

PCA Principal Component analysis

PCV Phenotypic Coefficient of Variation

SARC Sirinka Agricultural Research Center

SAS Statistical Analysis System

V
 TABLE of CONTENTS

CONTENTS PAGE

DEDICATION................................................................................................................................ I
STATEMENT OF THE AUTHOR ............................................................................................ II
BIOGRAPHICAL SKETCH ..................................................................................................... III
LIST of ACRONYMS and ABBREVIATIONS ....................................................................... V
LIST OF TABLES ...................................................................................................................... IX
LIST OF FIGURES ..................................................................................................................... X
LIST OF APPENDIXES ............................................................................................................ XI
1 INTRODUCTION...................................................................................................................... 1
2. LITERATURE REVIEW ........................................................................................................ 3
2.1. Origin and Taxonomy of Bread Wheat .................................................3
2.1.1. Reproductive and Floral Biology ...................................................................... 4
2.1.2. Pollination ......................................................................................................... 5
2. 2. Wheat Production .................................................................................5
2.2.1. Wheat Production in the World ........................................................................ 5
2.2.2. Wheat Production in Ethiopia ........................................................................... 6
2.2.3. Importance of Bread Wheat .............................................................................. 7
2. 2.4. Grain Yield in Bread Wheat ........................................................................... 8
2.3. History of Wheat Breeding and Genetics in Ethiopia ...........................8
2.3.1. Breeding Methods of Bread Wheat................................................................... 9
2. 4. Genetic Variability, Heritability in Broad sense and Genetic Advance
in Bread Wheat ..................................................................................9
2.4.1. Variability in Wheat .......................................................................................... 9
2.4.2. Genotypic and Phenotypic Coefficients of Variation ..................................... 12
2.4.3. Heritability in Broad sense (H2)...................................................................... 13
2.4.4. Genetic Advance ............................................................................................. 15
2. 5. Association among Characters ...........................................................16
2.5.1. Genotypic and Phenotypic Correlation Coefficients ...................................... 16

VI
 2.5.2. Path Coefficient Analysis ............................................................................... 17
2. 6. Principal Component Analysis...........................................................18
2. 7. Genetic Divergence (Distance) and Cluster Analysis ........................20
3. MATERIALS AND METHODS ........................................................................................... 22
3.1. Description of the Study Area .............................................................22
3.2. Planting Materials ...............................................................................22
3.3. Experimental Design, and Trial Management ....................................25
3.4. Data collected ......................................................................................25
3.4.1. Plant basis ....................................................................................................... 26
3.4.2. Plot basis ......................................................................................................... 26
3.5. Statistical Analysis ..............................................................................27
3.5.1. Analysis of Variance (ANOVA) ..................................................................... 27
3.5.2. Genotypic and Phenotypic Coefficients of Variation ..................................... 30
3.5.3. Genetic Advance (GA) ................................................................................... 30
3.5.3.2. Genetic advance as percent of mean ..........................................................30

3.5.4. Correlation Analysis ....................................................................................... 31

3.5.5. Path Coefficient Analysis ............................................................................... 31
3.6. Principal Component Analysis ............................................................32
3.7. Genetic Divergence and Cluster Analysis ..........................................32
4. RESULTS AND DISCUSSION ............................................................................................. 34
4.1. Analysis of Variance ...........................................................................34
4.2. Genetic Variability and Mean Performance of Genotypes .................35
4.3. Genotypic and Phenotypic Coefficients of Variation .........................36
4.4. Estimation of Heritability in Broad Sense ..........................................37
4.5. Estimates of Expected Genetic Advance (GA) ...................................39
4.6. Correlations Analysis of Quantitative Traits ......................................40
4.7. Path Coefficient Analysis ....................................................................46
4.8. Principal Components Analysis ..........................................................48

VII
 4.9. Genetic Divergence (Distance) Analysis ............................................50
4.10. Clustering of Genotypes ....................................................................51
4.11. Cluster Mean Analysis ......................................................................53
5. SUMMARY AND CONCLUSION ....................................................................................... 55
6. REFERENCES ........................................................................................................................ 57

VIII
 LIST OF TABLES

Tables Page

Table1. Harvested area, production and average yields for major wheat producing countries
during 2014 ...................................................................................................................... 6
Table 2. Description of bread wheat genotypes used in the study ............................................... 23
Table 3. Skeleton for individual location analysis of variance for simple lattice design ............. 28
Table 4. Analysis of variance in the case of a series of genotypes evaluated across environments
....................................................................................................................................................... 29
Table 5. Estimated values of mean squares C.V (%) and R-square (%) for 11 traits of 49 bread
wheat genotypes combined over across locations ......................................................... 35
Table 6. Estimates of range, means, genotypic and phenotypic variances, broad sense
heritability, genetic advance, and genetic advance as a percentage of mean for 11
characteristics of 49 bread wheat genotypes, combined across the locations ............. 40
Table 7. Genotypic correlation coefficient (rg) (upper diagonal) and phenotypic correlation
coefficient (rp) (below diagonal) of 11 traits of 49 bread wheat genotypes ................ 45
Table 8. Estimate of direct (bold face and diagonal) and indirect (off diagonal) effects at
genotypic level in 10 traits of 49 bread wheat genotypes ............................................ 48
Table 9. Vector loadings and percentage of explained variation by the first four PCs ................ 50
Table 10. Inter and intra (bold) cluster D2 values among six clusters in 49 bread wheat
genotypes ................................................................................................................... 51
Table 11. Distribution and grouping of 49 bread wheat genotypes into different diversity classes
of cluster membership based on D2 analysis ................................................................ 52
Table 12. Mean values of seven clusters for 11 characters of 49 bread wheat genotypes ........... 54

IX
 LIST OF FIGURES
Figure Page

Figure 1. Evolution of Cultivated Wheat ........................................................................... 4

Figure 2. Wheat Production, Area Cultivated and Yield in Ethiopia. ................................ 7

X
 LIST OF APPENDIXES

Appendix Tables Page

1. Mean values of 11 quantitative traits of 49 tested bread wheat genotypes ............................. 66

2. Estimated values of mean squares of ANOVA and genetic parameters of 49 bread wheat
genotypes for 12 traits tested at Jamma using simple lattice design ........................................ 68
3. Estimated values of mean squares of ANOVA and genetic parameters of 49 bread wheat
genotypes for 12 traits tested at Geregera using simple lattice design ..................................... 69

XI
Appendix Figures Page

1. Principal components plot of bread wheat genotypes based on 11 agronomic and phenotypic
traits. ......................................................................................................................................... 70
2. Tree diagram of genetic relationships among 49 bread wheat genotypes. ............................... 71

XII
GENETIC VARIABILITY AND CHARACTER ASSOCIATION IN BREAD
WHEAT (Triticum aestivum L.) GENOTYPES AT JAMMA AND GEREGERA,
ETHIOPIA
ABSTRACT

Continuous identification of the best genotypes that have wider genetic base, capable of
producing better yield under a wide range of agro-climatic conditions and stresses increases
production and productivity. Forty nine bread wheat genotypes were evaluated for 12 traits in
simple lattice design at Jamma and Geregera to determine the extent of genetic variation and
character association among grain yield and its related traits. Mean squares of the traits studied
showed statistically significant differences among the genotypes listed (P< 0.01), indicating the
presence of adequate variability. Maximum values of genotypic coefficient of variation were
recorded for spike length (8.66%), number of productive tillers (8.4%), number of grains per
spike (6.4%) and thousand seed weight (6.15%), whereas better value of phenotypic coefficient
of variation were recorded for productive tillers, grain yield, spike length and harvest index with
values of (13.3%, 11.35%, 10.3% and 9%), respectively. Heritability ranged from 29.1% for
grain yield to 82% for days to heading. Relatively high genetic advance as percent of mean was
obtained for spike length, productive tillers, number of grains per spike, thousand seed weight,
heading date and plant height with values of (14.9%, 10.6%, 10%, 10%, 9.7%, and 9%),
respectively. Grain yield had strong and positive genotypic correlation with harvest index
(0.731), biological yield (0.617), thousand seed weight (0.395), plant height (384) and
productive tillers (0.366). Path analysis indicated maximum positive direct effect obtained
between grain yield and harvest index (0.731) and also grain yield and biological yield (0.731).
The first five principal components, with eigenvalue greater than one, accounted for 80.4% of
the total variation. Based on the average linkage cluster analysis, the 49 genotypes were
classified into six clusters; indicating the genotypes were divergent. Thus, crossing program
between members of cluster I with cluster III, and cluster II with III, and IV could possibly
resulted in heterosis in the F1, and a great deal of variability in the F2. Plant selection based on
plant height, higher number of grains per spikes, thousand seed weight, biological yield and
higher harvest index will be most effective for future wheat yield improvement program.

Key words: Bread wheat genotypes, Character association, Genetic variability, Grain yield.

XIII
 1 INTRODUCTION
Bread wheat (Triticum aestivum L.) is a self-pollinating annual plant belonging to the family
Poaceae, and it is an allohexaploid species with three different genomes configuration (A, B, D)
of 42 chromosomes (2n = 6x = 42) (http://sundoc.). It is used as a domestic food consumption
and industrial crop (Gashaw et al., 2014), manufacture of flour for making bread and other home
made products (Negash et al., 2013). It is also traded food crop internationally, and an
emergency food in aid for developing countries (Hailu, 2011).

In 2014, 723.4 million ton wheat was produced from 222.3 million hectare (ha) with average
yield of 3.25 ton/ha worldwide including the main wheat producing regions of European Union,
China, India, Russian, United States, Canada, Australia, Pakistan and Ukraine (FAO, 2015,
USAD, 2015). Whereas the national share of wheat in cereal area was around 13.25% (1.66m ha)
in 2014, and share in production was 15.65% (4.23million metric ton) with average yield of 2.54
ton per ha (CSA, 2014/15). It was ranked third in total production as cereal behind maize and teff
and forth in area coverage after teff, maize and sorghum.

In Ethiopia, an effectively organized national wheat research program started since 1967 and
many varieties were released (Hailu, 1991). Attempts have been made so far to improve
production and productivity of bread wheat considering it importance as a food and industrial
crop, the ever increasing population, its high economic and nutritive value as well as its vast
acreage devoted to its cultivation. However, the average yield at production fields has remained
2.54 t/ha, which is low compared to the experimental yield of above 5 t/ha in the country (Hailu,
1991); the world average of 3.25 ton per ha (FAO, 2015, USAID, 2015); other leading wheat
producers in the world like Germany, France where average yields were 7.4 and 7.2 t/ha
respectively (Yao et al., 2012).

This is primarily due to the consequence of interaction between various abiotic and biotic
factors, shortage of high yielding genotype which are adapted to local conditions and stresses,
wide seasonal variability and environmental fluctuation, low amount of rainfall, and poor soil
moisture conservation (Hailu, 1991; Adem, 2013; Mideksa and Tadele, 2014). The important
biotic stresses include diseases, such as rusts and weed causes maximum yield losses of 30-50 %

1
and 29%, respectively (Samuel et al., 2014; Sramková et al., 2009). Drought also causes
maximum yield losses of 64%, which affects growth and development of plants through
alterations in metabolism and gene expression (Nezhadahmadi et al., 2013).

As a result, to alleviate those constraints, effective breeding program for grain yield
improvement to further yield increases, continue identification of best genotypes as donors of
various genes of agronomic importance as well as the development of superior cultivars depends
upon; various genetic and non-genetic factors, the amount of genetic variability present in the
plant population, and association of agro-morphological traits with grain yield (Ali et al., 2009).
Besides this genetic variability with the help of suitable parameters such as genetic coefficient of
variation, heritability estimates and genetic advance are necessary to start an efficient breeding
program in any crop plants including bread wheat (Ali et al., 2009; Fellahi et al., 213; Kumar et
al., 2014). With continuous phenotypic traits, in most cases the alleles that are present in the
population or even the loci affect the trait are unknown, so we need to use the statistical
measures of mean and variance (and later covariance) to characterize populations.

The characterization of this variability in a population is relevant since genetic diversity within
population and within species determines the rates of adaptive evolution and the extent of
response in bread wheat improvement. Knowledge of genetic diversity and the genetic
relationship between genotypes is equally important consideration for efficient rationalization
and utilization of germplasm resources. Information on genetic diversity is also needed for the
optimal design of plant breeding programmes, influencing the choice of genotypes to cross for
the development of new populations (http://sundoc.) Therefore, this study was conducted with
the following objectives:

General objective

To estimate the extent of genetic variability, association of grain yield with other characters as
well as direct and indirect effects of yield attributing traits on bread wheat grain yield.

Specific objectives

i. To determine variability, heritability and genetic advance

ii. To estimate the correlation and path coefficients among traits studied

2
 2. LITERATURE REVIEW

2.1. Origin and Taxonomy of Bread Wheat

The probable origin of the genus Triticum is found in Asia, in the area known as the Fertile
Crescent, and parts of Africa, in the area that stretches from Syria to Kashmir and southwards to
Ethiopia (Singh and Kota, 2007; Sramkova et al., 2009). The central Asia, Near East,
Mediterranean and Ethiopian regions are origins and centers of diversity of wheat species, and
had distributed to India, Great Britain, Ireland and Spain (Sramkova et al., 2009).Taxonomically
it belongs to the family Poaceae(grasses), tribe Triticeae, genus Triticum and species aestivum
(Acquaah, 2007).

The genetic origin of wheat is a classic example how closely related species combine in nature to
form a polyploidy series (Singh and Kota, 2007). The genome analysis, the determination of
evolutionary relationships on the basis of chromosome pairing in hybrids indicated that
allopolyploidy was involved and that wheat evolution followed a system of diploid divergence
and polyploid convergence (http://www.oecd.org/ehs). All the species of wheat are grouped into
three groups: diploid, tetraploid and hexaploid that form a polyploid series with chromosome
numbers, a turning point in triticum classification, 14 (n=7), 28 (n=14) and 42 (n=21)
(Sramková, et al., 2009), respectively.

Evidences indicate that the hexaploid wheat (AABBDD) is believed to have arisen when
genomes of diploid (2n = 14, AA) forms of T. monococcum(a) were naturally pollinated by weed
species (2n = 14, BB). The subsequent genome duplication of hybrids by natural polyploidy gave
rise to several wild and cultivated tetraploid species (2n = 28, AABB) like T. dicoccum(b) and T.
durum; again, the natural pollination of the tetraploid T. dicoccum(b) by Aegilops squarrosa
L.(Ttiticum tauschii) (2n =14, DD) gave rise to the hexaploid (2n = 42, AABBDD) species (c)
(Singh et al, 2007; Sramková et al., 2009; Velu and Singh, 2013).

The haploid DNA content of hexaploid wheat is approximately 1.7 x 1010base pair (bp) genome
size resulted from polyploidy and extensive duplications, such that over 80% of the genome
consists of repetitive DNA sequences. This base pair is about 100 times larger than the
Arabidopsis genome, 40 times that of rice and about 6 times that of maize(http://sundoc.).

3
 .
Figure 1. Evolution of Cultivated Wheat
Source: Sramkova et al. (2009).

2.1.1. Reproductive and Floral Biology

According to Acquaah (2007), wheat has a determinate, composite spike inflorescence. Each
spike bears 10–30 spikelets, which are borne singly at nodes on alternate sides of a zigzag rachis.
A spike may be awnless, awnleted, or awned. A spikelet consists of 1–5 flowers (or florets)
attached alternatively to opposite sides of the rachilla (central axis). A spikelet is subtended by a
pair of empty bracts and glumes (Singh and Kota, 2007).

A floret consists of a lemma and pale, which enclose these stamens and a pistil, plus two
lodicules that regulate the opening of the flowers and anthers. Wheat flowers bloom under
temperatures of 13–25°C. The flowering is usually diurnal, the highest peak occurring in the
morning, and a lower peak in the afternoon. Blooming begins in the spikelets located above the
middle of the spike and proceeds both upward and downward. It takes about 2–3 days for a

4
wheat spike to complete blooming, after the appearance of the first anthers. The flowering period
may last from 14 to 21 days (Singh and Kota, 2007).

2.1.2. Pollination
Wheat is predominantly self-pollinated with about 1–4% natural cross-pollination (Singh and
Kota, 2007). Pollen shed usually starts inside the floret, but about 80% of anther dehiscence
occurs outside the floret. The primary and secondary florets produce larger and more viable
pollen grains than other florets. Wheat pollen remains viable for up to about 30 minutes after
shedding. Once pollinated, the pollen tube growth starts within 15–60 minutes. Even though the
stigma remains receptive for up to 13 days, it is most receptive within 3 days of anthesis.

2. 2. Wheat Production

2.2.1. Wheat Production in the World

Wheat is the staple food of the 1/3rd of the world’s population. Approximately 25 % of global
agricultural land is utilized for wheat cultivation, making it the largest food crop worldwide in
terms of area (Velu and Singh, (2013), and the productivity is increasing at less than 1%
annually, while the annual productivity must increase at 2 % annually to meet the global
demand. Because of its high economic and nutritive value, vast acreage devoted to its cultivation
and its association with some of the earliest and most important civilizations of the world, wheat
is known as the queen of cereals. It is also the first strategic, and the world’s leading cereal grain
(Chhibber et. al., 2014; Saleem, et al, 2015). It is mostly a temperate crop, while it grows in a
wide range of agro-climatic regions under several production systems worldwide (between
latitudes 30° and 60° north and south) at altitudes from sea level to 3000 m.a.s.l with an optimum
growing temperature of 25°C (Sramkova et al., 2009).

Globally, wheat is the leading source of vegetable protein in human food, having a higher protein
content than other major cereals, maize (corn) or rice. In terms of total production used for food,
it is currently second to rice as the main human food crop and ahead of maize, after allowing for
maize's more extensive use in animal feeds (Hailu, 2011).

Wheat requirement is increasing continuously due to ever increasing population of the world
(Waqas et al., 2014), while the yield is generally insufficient to fulfill the domestic requirements

5
 (GAIN, 2014), which calls for improved and high-yielding varieties to be developed by plant
breeders. Therefore, it is necessary to develop new wheat cultivars, as well as to enhance the
existing ones, that are genetically more stable, having wider genetic base, capable of producing
better yield under a wide range of agro-climatic conditions and stresses to improve the yield
potential and enhance the grain yield (Ali et al., 2008; Laghari et al., 2010; El-Mohsen et al.,
2012; Farshadfar et al., 2013; Fellahi et al, 2013; Kumar et al, 2014).

Table1. Harvested area, production and average yields for major wheat producing
countries during 2014

SN Country Area harvested (million ha) Production (million ton) Yield (Metric ton/ha)

Worldwide
1 European Union 26.8 155.6 5.81
2 China 24.1 126.2 5.23
3 India 30.6 95.80 3.13
4 Russia 23.4 59.0 2.23
5 U.S.A 18.8 55.1 2.94
6 Canada 9.46 29.3 3.10
7 Australia 13,8 26.50 1.74
8 Pakistan - 25.3 -
9 Ukrine 6.3 24.5 3.89
In Africa
1 Egypt 8.8
2 Morocco 5.1
3 Ethiopia 1.665 4.23 2.54
Total Worldwide 222.288 723.384 3.25
Sources: CSA (2014/2015), FAO (2015), USDA (2015).

2.2.2. Wheat Production in Ethiopia

In Ethiopia, it contributes a major part in achieving the millennium goal of the country, food
grain self-sufficiency (Mathewos and Tewodros, 2012; Tewodros et al., 2014). The most suitable

6
areas fall between 1800-2800 m.a.s.l with average temperature and rain fall ranges from 15-25
0
C and 400-1200 mm, respectively (Adem, 2013). Almost all wheat production (more than 99%)
comes from four major wheat producing regions; Oromia, Amhara, SNNPR and Tigray regions
(CSA, 2013). From 1995/96 to 2012/13 wheat production area and yield increased by double
from 0.8 million ton to 1.6 million ton, and from 1.2 t/ha to 2.1 t/ha, respectively (CSA, 2013).

Figure 2. Wheat Production, Area Cultivated and Yield in Ethiopia.

Source: FAOSTAT (2014) cited in Gashaw et al. (2014).

2.2.3. Importance of Bread Wheat

Wheat is one of the major staple crops in the country in terms of both production and
consumption (Gashaw et al., 2014). It is used for the manufacture of flour for bread, biscuits and
industrial products (Mathewos and Tewodros, 2012; Negash et al., 2013). Wheat grain is a staple
food used to make flour for leavened, flat and steamed breads, biscuits, cookies, cakes, breakfast
cereal, couscous and for fermentation to make beer, other alcoholic beverages and biofuel
(https://en.wikipedia.org/wi). Traditionally the crop is used for making dabo, dabokolo, ganfo,
kinche and other types of foods (Mathewos and Tewodros, 2012; Negash et al., 2013). The straw
is good source for animal feed and is also used for thatching roofs (Mathewos and Tewodros,
2012; GAIN, 2014). The basic ingredients of bread wheat grain are carbohydrate (70%), water
(12%), protein (12%), vitamins and minerals (2%), lipid (2%) and crude fiber (2%) (White and
Edwards, 2008).

7
2. 2.4. Grain Yield in Bread Wheat

The wheat grain is caryopsis, a small dry, indehiscent; one seeded fruit with a thin pericarp
consisting of a germ or embryo and an endosperm (Singh and Kota, 2007), as well as seed is the
reproductive unit and the end-use product. Grain yield in wheat is a complexly inherited trait of
low to moderate heritability and is strongly influenced by the environmental conditions (Velu
and Prakash, 2013). A wheat grain can be broadly divided into three components: seed coat and
aleurone layer (or bran) (14%), endosperm (83%), embryo (germ) (3%). Grain development is
the period from flowering to physiological maturity when fertilized florets fill and ripen to form
grain. The wheat grain has three growth stages; grain enlargement, grain fill and physiological
maturity (White and Edwards, 2008).

In many crops, a variation of genotypes in time to reproductive stage is a source of a

combination of genetic and environmental constraints and requires appropriate consideration. In
general, unfavorable conditions in time to reproductive stage differently affects productivity and
growing of commercial cultivars in production areas. Thus, genotypes least effected from
changed environmental conditions especially in reproductive stage can remain present in yield
performance (Nezhadahmadi et al., 2013).

To attain maximum yield, it is important to achieve a balance between biomass and resources.
The inputs that can be managed are: plant population, fertilizer, sowing date, diseases and
insects, row spacing, surface stubble. Yield is determined by four components: number of
heads/m2, number of spikelet per head, number of grains per spikelet, weight per grain (White
and Edwards, 2008). Yield potential is generally assessed through grain yield and yield
components, which themselves are complex characters and are considered to be the cumulative
result of different physiologic processes (Ali et al., 2009).

2.3. History of Wheat Breeding and Genetics in Ethiopia

In fact research on wheat in Ethiopia has been active for more than six decades, it has passed
through different phases and has never fully satisfied the needs of farmers in the different wheat
production systems. Wheat research in Ethiopia prior to 1930, dealt mainly with germplasm
collection, identification and characterization. From 1930-52, introduction, hybridization and

8
selection began, culminating in the release of Kenya 1 and 5. This work continued at Debre Zeit
and other stations during the period 1953-66 when 6 cultivars were released. The organized
national wheat improvement program has been started most effectively from 1967 and from
1967 up to 1990, thirty improved wheat varieties have been released (Hailu, 1991).

2.3.1. Breeding Methods of Bread Wheat

The choice and key success in plant breeding work mainly depends on knowledge of the crops;
mode of reproduction (sexual reproduction, asexual reproduction), system of pollination (self-
pollinated, cross-pollinated), gene action of the desired characters (additive, dominance,
epistasis), sufficient genetic variations; types of variation (non-heritable and heritable variation)
and kinds of variation (qualitative and quantitative variation), appropriate method of selection
(related to gene action of the characters, types of variety going to develop), selection criteria
being developed (single character being bred and multiple selection criteria) and breeding
objective of crop species (Acquaah, 2007). The main structures of the wheat plant are the
coleoptiles, leaves, tillers, stem, roots and head (White and Edwards, 2008), and growth is
determined by number of tillers/plant, root and shoot lengths and fresh and dry weights (Shahzad
et al., 2012).

Being self- pollinated crop, the basic methods of wheat improvement include pure line, pedigree,
bulk, single-seed descent and back cross method (Baenziger and DePauw, 2009). The first phase
for development of improved wheat genotypes is the adoption of pure line selection from
indigenous landraces and then introduction of improved exotic types. Later hybridization
program involving inter crossing of systematically selected genotypes in a system of single,
double or complex multiple crossing schemes followed by various forms of pedigree selection.

2. 4. Genetic Variability, Heritability in Broad sense and Genetic Advance in

Bread Wheat

2.4.1. Variability in Wheat

Variation is differences between parents and their offspring or among individuals in a

population. It is an important aspect of breeding, for if all organism look alike there will be no

9
bases for breeding work. Variation in a population or among a group of individual therefore is
important to the breeder (http://www.unaab.edu.ng).

Genetic variability is a measure of the tendency of individual genotypes in a population to vary

from one another. Variability is different from genetic diversity, which is the amount of variation
seen in a particular population. The variability of a trait describes how much that trait tends to
vary in response to environmental and genetic influences. Genetic variability in a population is
important for biodiversity, because without variability, it becomes difficult for a population to
adapt to environmental changes and therefore makes it more prone to extinction
(http://www.unaab.edu.ng).

Variability is an important factor in evolution as it affects an individual's response to

environmental stress and thus can lead to differential survival of organisms within a population
due to natural selection of the fit variants. Variability results due to differences either in the
genetic constitution of the individuals of a population or in the environment in which they have
grown. The quantitative measurement of individual character provides the basis for an
interpretation of different variability parameters. The phenotypic variability which is observable
includes both genotypic and environmental variation and changes under different environmental
conditions (Farshadfar et al., 2013).

Thus, separating the total variation into heritable and non-heritable components with the help of
genetic parameters i.e. genotypic and phenotypic co-efficient of variation, heritability and
genetic gain is necessary (Kahrizi et al., 2010; Farshadfar et al., 2013). Often, it is not feasible to
determine the number of genes affecting a particular trait, and the individual effects of genes on
the phenotype. The extent to which variation in yield components are responsible for differences
in yield among various genotypes, it must be borne in mind that overall variability depends on
heritable and non-heritable components (Tabbal, 2012).

Biological variation exist in the plant population are phenotypic, genotypic and environmental,
but a primary step in wheat cultivar improvement is to generate heritable genetic variation,
which remains unaltered by environmental conditions and is more useful to a plant breeder for
exploitation in selection or hybridization (Farshadfar et al., 2013; Kumar et al., 2014). The
development of an effective plant breeding program and the efficiency of selection mainly

10
depends upon the magnitude of genetic variability existed in plant material under study, because
it is pre-requisite for finding nature and extent of association among various yield and yield
components (Ali et al., 2008; Laghari et al., 2010; El-Mohsen et al., 2012; Farshadfar et al.,
2013; Fellahi et al, 2013; Kumar et al, 2014; Adhiena et al., 2016).

Many studies have investigated on the extent of genetic variability available in bread wheat.
Genetic variability studies conducted to investigate diversity for various traits in bread wheat
showed the existence of wide trait diversity that would respond positively to selection
(Moghaddam et al., 997; Laghari et al., 2010; Mathewos and Tewodros, 2012; El-Mohsen et al,
2012; Awale et al., 2013; Karim et al., 2013; Farshadfar et al., 2013; Fellahi et al., 2013; Kumar
et al., 2014; Adhiena et al., 2016).

Moghaddam et al. (1997) conducted an experiment to estimate genetic variation and heritability
for 13 developmental and quantitative characters in fifty-three pure lines of bread wheat and
reported highly significant variations among the materials for all characters like days to heading
ranged from (102–129), plant height (cm) 81–118), number of tillers per plant (5–11), number
of spikes per plant (4–8), main spike length (7–12 cm), number of grains per spike (20–54),
1000-grain weight (21–47g), shoot biomass per plant (15–29g), grain yield per plant (3–10g),
straw biomass per plant (7–20g ), harvest index (20–52(%)).

Ali et al. (2008) conducted the experiment to evaluate variability parameters, correlations and
path coefficients in seventy bread wheat genotypes for eight metric traits i.e., plant height, which
ranged from (64.57–120.17 cm), number of productive tillers per plant (5.33–24), number of
spikelets per spike (8.50–25.67), spike length (7.47–17 cm), number of grains per spike (22–
85.67), fertility % (80.15–97.83), 1000 grain weight (32.3–56.9 g), yield per plant (5.67–36.45
g). Thus he concluded that significant genotypic differences were observed for all the traits
studied indicating considerable amount of variation among genotypes for each character, which
provided a good opportunity for yield improvement.

Mohibullah et al. (2011) conducted a study for five quantitative characters in hundred breed
wheat germplasm to test the variation with correlation and reported a wide range of variation and
highly significant variation for characters studied like spike length (6.9 -22.4 cm), number of
spikelets per spike (10.5- 31.8), grain yield per plan (1.35 -4.6), 1000-grain weight (16.8- 46.2),

11
grain yield (2701- 5185 kg /ha). Mathewos and Tewodros (2012) found large variation for
morphological traits such as plant height ranged from (72.8-120.6 cm), spike length (3.9-9.95
cm), number of seeds per spike (25.5-70.2), number of days to mature (83-167) and yield (0.46
to 12.4 t/ha) over locations.

El-Mohsen et al. (2012) studied eight quantitative characters of ten wheat varieties and reported
a wide range of variation and highly significant variation for characters studied like days to
(50%) heading (80.1-95.44), plant height (70.6-122.3), number of tillers per plant (5.89-20.36),
spike length (8.65-14.22), number of spikelet’s per spike (15.4-25.7), number of grains per spike
(33.56-71.27), thousand grain weight (35.27-58.42g) and grain yield per plant (10.22-39.25g).

Studies of Awale et al. (2013) conducted to estimate the extent of genetic variability and traits
association in bread wheat genotypes , reported a wide range of variation for characters studied
like days to heading (32 to 71 days), grain filling period (11 to 66 days), days to maturity (59 to
105 days), plant height (35 to 68 cm), number of tillers per plant (1 to 8), spike length (1 to 13
cm), number of spikelet’s per spike (2.2 to 17), number of grains per spike (31.7 to 59), 1000-
grain weight (10.6 to 54g) and grain yield per plot (12.1-48.2 qt/ha) indicating good opportunity
for grain yield improvement.

Studies conducted at Ofla district, North Ethiopian, in 2014 with the objective of estimating
nature and magnitude of variation existing in twenty six bread wheat genotypes (Adhiena et al.,
2016), reported highly significant variations among genotypes ranged for days to heading (49.33-
66), days to maturity(102.7-129.7), grain filling period (47.67-66.67), number of tillers (1.53-
3.27), spike length (5.87-8.87), number of kernels per spike (34.27-46.7), 1000-kernel weight
(68.53-95.20), biological yield (6.46-16.17), grain yield (2.96-6.35 qt/ha).

2.4.2. Genotypic and Phenotypic Coefficients of Variation

Coefficients of variation measure the magnitude of variability present in a population. The

success or failure in breeding program largely depends on the extent of variability in the base
population which is measured by different population parameters including genotypic and
phenotypic coefficients of variation (Usmani et al., 2014).

12
Several researchers have estimated the genotypic and phenotypic coefficient of variation
between different yield attributing characters and their effects on yield in bread wheat. The high
phenotypic coefficient of variation (PCV) and genotypic coefficient of variation (GCV) indicate
the high variability of characters in the germplasm that selection may be effective based on these
traits and their phenotypic expression would be good indication of the genotypic potential (Ali et
al., 2008; El-Mohsen et al., 2012; Awale et al., 2013; Kumar et al., 2014; Adhiena et al., 2016).
Many workers demonstrated higher phenotypic coefficient of variation than the corresponding
genotypic coefficient of variation which indicates less effect of environment on the expression of
characters studied (Moghaddam et al., 1997; Kashif et al., 2004; Ali et al., 2008; Atta et al.,
2008; Laghari et al., 2010; Kalimullah et al., 2012; Farshadfar et al., 2014).

On an average, higher magnitude of GCV and PCV were recorded for grain yield per plant,
harvest index, tillers per plant, spike length, number of kernels per spike, biomass yield and test
weight suggesting sufficient variability and thus scope for genetic improvement through
selection for these traits (Moghaddam et al., 1997; Kashif et al., 2004; Ali et al., 2008; Atta et
al., 2008; Laghari et al., 2010; Kalimullah et al., 2012; Farshadfar et al., 2014).

2.4.3. Heritability in Broad sense (H2)

Quantitative traits are influenced by genetic and environmental factors. The gross variation in a
population is the result of the combination of genotypic and environmental effects. Most of this
dissimilarity caused by the genotype is called heritability (Kumar et al., 2014). The genotypic
component being the heritable part of the total variability, its magnitude on yield and its
component characters affects the selection strategies to be adopted by the breeders (Ahmed et al.,
2007). A useful thing for breeders to know is, for any trait of interest, how much of the
phenotypic variability of that trait is due to genetic variance, and how much is due to non-genetic
environmental factors (http://www.unaab.edu.ng). Complex traits show a low heritability
because their expression is highly influenced by environmental factors, i.e. the conditions in
which the genotype is grown. First, there is no one heritability for given trait in a given species,
because heritability can and often does differ among populations and among environments. It can
differ among populations because additive variance (VA) depends on allele frequencies
(http://www.unaab.edu.ng).

13
The heritability (H2) of a character, which is the proportion of phenotypic variance due to
variance in genes (H2 = VG/VP x100), could vary from 0-100% (Baenziger and DePauw, 2009).
The heritability value of 0% indicates that genes do not contribute at all to phenotypic individual
differences, and the heritability value of 100% indicates genes are the only reason for individual
differences (heritability/heritability.intro.html). Heritability plays an important role in deciding
the suitability and strategy for selection of a trait. Heritability estimates show the efficiency in
which selection of genotypes can be based on phenotypic performance of quantitative traits
(Acquaah, 2007). Heritability estimates under stress conditions were found to be lower than
under controlled conditions which indicated that heritability is not constant and varies with
changes in environment (Shahzad et al., 2012). Characteristics of tolerant varieties grown in
stress environments relative to their performance without stress(Hailu,1991) are; the maturity
period is reduced by 20 %, plant height is reduced by less than 30%, harvest index ranges
between 0.3 - 0.5, kernel weight exhibits a low variance, and grain yield is reduced by less than
50%.

Studies have been conducted to estimate heritability in bread wheat and the higher the
heritability estimates, the simpler are the selection procedures (Moghaddam et al., 1997; Ali et
al., 2008; Laghari et al., 2010; Kalimullah et al., 2012; Awale et al., (2013); Karim et a., 2013;
Farshadfar et al., 2013; Fellahi et al., 2013; Kumar et al., 2014; Tewodros et al., 2014; Adhiena
et al., 2016). According to Kumar et al. (2014), traits closely associated with yield should be
more heritable than per se to serve as better indicators of the genetic yield potential of a line.

Estimating of heritability was conducted in grain yield of bread wheat and high heritability were
reported (Moghaddam et al.,1997; Kashif et al., 2004; Ali et al., 2008; Kalimullah et al., 2012;
Awale et al., 2013; Farshadfar et al., 2014; Kumar et al., 2014). However, Laghari et al. (2010)
reported moderate heritability in grain yield. Moreover, Mohammadi et al. (2011), Mollasadeghi
et al. (2012); Tesfaye et al. (2014) and Adhiena et al. (2016) reported low heritability in grain
yield such as (7.4%), (12.27%), 19% and 25%, respectively, indicating the limited scope of
improvement of this trait through selection.

Estimating of heritability was made in characters like heading date, maturity date, number of
tillers per plant, plant height and high heritability were recorded and reported (Moghaddam et

14
al., 1997; Laghari et al., 2010; Awale et al., 2013; Farshadfar et al., 2014; Kumar et al., 2014;
Adhiena et al., 2016). However, Ali et al., (2008) and Mollasadeghi et al. (2012) reported
moderate heritability in number of productive tillers, and Adhiena et al. (2016) reported low
heritability in tillers per plant (4.45%). Laghari et al. (2010) also reported low heritability in
plant height.

Moderate high to high heritability for traits like spike length, number of spike lets per spike,
number of grains per spike, 1000-grain weight due to smaller phenotypic variances were reported
(Moghaddam et al., 1997; Kashif et al., 2004; Ali et al., 2008; Atta et al., 2008; Laghari et al.,
2010; Kalimullah et al., 2012; Awale et al., 2013; Karim et al., 2013; Farshadfar et al., 2014;
Adhiena et al., 2016). However, Laghari et al. (2010) reported moderate heritability for number
of grains per spike, and Awale et al. (2013) reported moderate heritability for number of
spikelets per spike (57.40%) and low heritability for spike length (23.48%).

High heritability was observed in harvest index (Moghaddam et al., 1997; Kumar et al., 2014)
and biological yield (Kumar et al., 2014). On the other hand, Adhiena et al. (2016) reported
moderate heritability and low heritability for harvest index and biological yield, respectively.

2.4.4. Genetic Advance

Genetic advance (GA) is the superiority of selected individuals over the base (original)
population. Genetic advance under selection is a genotypic value, which depends on three things
(Allard, 1960): genetic variability, heritability or masking effect of non-genetic variability on the
genetic variability and the selection intensity applied. Genetic progress would increase with
increase in the variance. Level of improvement in one or more measured traits as compared to
natural or unimproved populations usually expressed as a percentage, and usually improvement
(GA) is determined by heritability of the trait (h2), selection and phenotypic standard deviation.
Because of the cyclic nature of a breeding program, the majority of parents in any given cycle
are represented by the best lines selected from the previous cycle (Kumar et al., 2014).

Moghaddam et al. (1997), Laghari et al. (2010) and Awale et al. (2013) reported high genetic
advance as percentage of mean in bread wheat for characters like number of tillers per plant,
plant height, spike length, number of spikelets per spike, number of grains per spike, 1000-seed

15
weight, harvest index, biological yield and grain yield, and also low genetic advance as a
percentage of mean for heading date and maturity date were reported. However, Moghaddam et
al. (1997), Laghari et al. (2010) and Awale et al. (2013) reported high genetic advance as a
percentage of mean for heading date and maturity date. On the other hand, Adhiena et al. (2016)
computed moderate genetic advance as percent of mean for most of traits studied except for
number of fertile tillers per plant, which refers to improvement of these traits in genotypic value
for the new population compared with the base population with one cycle of selection is not
rewarding.

2. 5. Association among Characters

2.5.1. Genotypic and Phenotypic Correlation Coefficients

A correlation coefficient gives a numerical summary of the degree of association between two
variables for example, to what degree do high values of one variable go with high values of the
other one? Correlation coefficient vary from -1 to +1, with positive values indicating an
increasing relationship and negative values indicating a decreasing relationship.

Correlation coefficient is an important statistical method, which can help breeders in selection
for higher yields. The correlation coefficient, the degree of association between two characters,
measures the magnitude and direction of mutual relationship between various plant characters
and helpful in determining the component characters of a complex trait, like yield (Mohammadi
et al., 2012; Laghari et al., 2010). The phenotypic and genotypic correlation coefficients are
measures of the degree of closeness of the linear relationship between pairs of variables and also
a value to indicate the degree to which various morpho-physiological characters are associated
with economic productivity (El-Mohsen et al., 2012).

The existence of correlation between a complex trait and its components is an indication of gene
association or pleiotropism. Correlations in phenotype may be due to genetic or environmental
causes and may be positive or negative. Genetic causes may be due to pleiotropy, linkage,
gametic phase disequilibrium. At genetic level, a positive correlation occurs due to coupling
phase of linkage and negative correlation occurs due to repulsive phase of linkage of genes
controlling two different traits.

16
Correlation coefficients (r) were estimated to study the relationships among the traits (Usmani et
al., 2014) ranged from -1 to +1 and r value of -1 or +1 indicates perfect correlation; values close
to -1 indicate high negative correlation. In opposition, values close to +1 indicate high positive
correlation. If there is no linear association between variables, the correlation is zero (Fellahi et
al., 2013). Statistical analysis showed that genotypic correlation coefficients were higher than the
corresponding phenotypic correlation coefficients in most of the traits which reflect the influence
of environment on the expression of traits (Ali et al., 2008; Laghari et al., 2010; El-Mohsen et
al., 2012; Kalimullah et al., 2012; Farshadfar et al., 2014).

Results of positive associations of grain yield per plant with number of tillers per plant, number
of spikelet’s per spike, spike length, number of grains per spike and 1000-grain weight at
genotypic and phenotypic levels were reported, but days to 50% heading and plant height
contributed negatively towards grain yield at both levels (Kashif et al., 2004; Ali et al., 2008;
Khan et al., 2010; El-Mohsen et al., 2012; Irfaq et al., 2012; Awale et al., 2013; Gelalch and
Hanchinal, (2013). However, Kashif et al., (2004) reported positive association between plant
height and grain yield.

2.5.2. Path Coefficient Analysis

In plant breeding program, direct selection for yield as such could be misleading. A successful
selection depends upon the information on the genetic variability and association of morpho-
agronomic traits with grain yield (Ali et al., 2008). The correlation coefficient may not give
sufficient information about the relationship between different variables as much as statistical
multivariate methods give (Fellahi et al., 2013). Therefore, correlation studies along with path
analysis provide a better understanding and an exact picture of the association of different
characters with grain yield (Ali et al., 2008). Path coefficient analysis provides an effective way
of finding out direct and indirect sources of correlations, using genotypic correlations of different
plant attributes (Kashif et al., 2004).

Path coefficient analysis is an efficient statistical technique especially designed to determine the
direct effect of one character on another character and permits the separation of a correlation
coefficient in to components of direct and indirect influences for a set of a priori cause and effect

17
interrelationships (Ali et al., 2008; Fellahi et al., 2013). Path coefficient analysis is a reliable
statistical technique, which provides means to quantify the interrelationship of different yield
components and indicate whether the influence is directly reflected in the yield or take some
other path ways to produce an effect. Grain yield per plant was selected as resultant variable and
plant height, flag leaf area, fertile tillers per plant, spike length, spikelets per spike, grains per
spike and 1000-grain weight as casual variables (Kashif et al., 2004).

Studies have been conducted to estimate direct and indirect effects of different traits on grain
yield in bread wheat. Khan et al. (2010) reported the highest direct effect of grains per spike on
grain yield followed by spike length and days to maturity whereas 1000-grain weight and plant
height had negative direct effect on the same parameter, and characters such as days to maturity
and grains per spike exerted positive direct effect along with positive genotypic correlation on
grain yield.

In a study conducted by Ali et al. (2008), number of grains per spike exhibited the highest
positive direct effect followed by number of productive tillers per plant and 1000-grain weight.
Furthermore, El-Mohsen et al. (2012) reported maximum positive direct effects of number of
grains per spike, followed by number of tillers per plant and 1000-grain weight on grain yield per
plant.

Path coefficient analysis study in bread wheat displayed maximum positive direct effect on grain
yield per plot mostly exerted by days to heading, grain filling period, number of tillers per plant
and grains per spike on grain yield per plot (Awale et al., 2013). Gelalcha and Hanchinal (2013)
also reported biomass, harvest index and plant height imparted significant direct influence on
grain yield in bread wheat.

2. 6. Principal Component Analysis

Principal component analysis (PCA) is defined as a method of data reduction to clarify the
relationships between two or more characters and to divide the total variance of the original
characters into a limited number of uncorrelated new variables (Fellahi et al., 2013). This will
allow visualization of the differences among the individuals and identify possible groups. The
reduction is achieved by linear transformation of the original variables into a new set of

18
uncorrelated variables known as principal components (PCs). The first step in PCA is to
calculate Eigen values, which define the amount of total variation that is displayed on the PC
axis.

Azeb (2016) noted the first PC generalizes most of the variability present in the original data relative to
all remaining PCs and a study conducted at Atsbi, Ofla and Quiha environments revealed that four
principal components PC1, PC2, PC3 and PC4 with eigenvalues 3.87, 2.87, 1.26 and 1.04, respectively
have accounted for 82.16% of the total variation among genotypes for the 11 quantitative traits considered
at Atsbi. This holds true for Ofla site, where the first three principal components PC1, PC2 and PC3 with
eigenvalues of 4.08, 2.07 and 1.47 resulted in 69.27% total variation and, the first three principal
components PC1, PC2 and PC3 with eigenvalues 4.64, 1.93 and 1.35, respectively, have accounted for
72.04% of the total variation where the first two principal components PC1 and PC2 contributed values of
42.20% and 17.58%, respectively to the total variation at Quiha environment..

Different levels of diversity were observed in different accessions/populations on the basis of

agro-morphological traits. The Eigen values are often used to determine how many factors to
retain and the sum of the Eigen values is usually equal to the number of variables (Fellahi et al.,
2013). According to the results of Fellahi et al. (2013), the estimated wheat variables had
grouped into three principal component factors with Eigen values more than one which all
together explained 77.44% of total variability.

Mideksa et al. (2014) reported principal component analysis explained better the variation
among varieties and land races which showed that the first four components with Eigen values
more than one accounted 74% (22, 21, 18 and 12%, respectively) plant height, ear shape, days to
maturity, grain yield, biological yield, ear color and grain color being the most important factors
in contributing to the total variability. The study of Awale and Sentayehu (2013) demonstrated
that characters having relatively higher value in the first principal component like number of
tillers per plant, grain yield per plot, grains per plot which accounted 27.9% from the total
variation (91.87%) of the six principal components had more contribution to the total variation
and they were the ones that most differentiated the clusters.

Daniel et al. (2011) assessed the genetic diversity for yield and yield related traits in 49 bread
wheat genotypes for 17 characters and showed wide variability for the components studied.
Accordingly, the result of the principal components analysis revealed that nine principal

19
components accounted nearly 80 % of the total variation. Out of the total principal components
retained, PC1, PC3, PC8 and PC 4 with values of 18.71 %, 9.68 %, 9.22 % and 8.15 %,
respectively, contributed more to the total variation. Hence, the first principal component had
high positive component loading from protein content, sedimentation volume and wet gluten
content; and high negative loading from grain yield, biomass yield and starch content contributed
more to the diversity and they were the ones that most differentiated the clusters. More over the
major contributing characters for the diversity in the third principal component (PC3) were days
to maturity, harvest index and days to heading; whereas grain filling period and days to heading
in principal component four (PC4); and biological yield, days to heading, grain yield and starch
content in principal component eight (PC8).

2. 7. Genetic Divergence (Distance) and Cluster Analysis

Genetic divergence is one of the important biometrical techniques used for estimating the extent of
diversity existed among selected genotypes present in a population (Mahalanobis, 1936). The pattern
and level of genetic diversity in a given crop gene pool can be measured in terms of genetic distances
which is the extent of gene differences between cultivars as measured by allele frequencies at sample loci.

Precise information on the nature and degree of genetic diversity helps the plant breeder in
choosing the diverse parents for purposeful hybridization since it is necessary that the varieties
should be genetically divergent especially for quantitative characters that contribute towards yield
(Daniel et al., 2011). The analysis of genetic diversity through the cluster analysis of cluster
diagram (dendrogram) based on different dissimilarity/similarity measures using various
clustering algorithms categorize genotypes in to different clusters at a certain percentage of
dissimilarity/similarity level. Knowledge of genetic diversity and relationships among elite breeding
materials is important for the improvement of crop plants. Genetic diversity is different
from genetic variability in a way that the former measures the number of the actual variation
of species in a population whereas the latter measures how much the trait or the genotype will
tend to vary. Genetic diversity refers to any variation in the nucleotides, genes, chromosomes, or
whole genomes of organisms (http://www.unaab.edu.ng).

Cluster analysis is a multivariate method or technique, which aims to classify a sample

genotypes based on a set of measured variables into a number of different groups such that

20
similar genotypes are placed in the same group and arranging variables into different clusters to
find the clusters that their cases within are more similar and correlated to one another comparing
to other clusters (Fellahi et al., 2013).

Clustering is also defined as the process of organizing genotypes into homogeneous groups and
is performed to study the patterns of groupings of genotypes whose members are similar in some
way (Chahal et al., 2002). It operate on a matrix of dissimilarity (or similarity) indexes for all
possible pairs of genotypes depending on which is being clustered (Ghaderi et al., 1980).

In a study conducted by Ali et al. (2008), cluster analysis grouped 70 wheat genotypes into 4
different clusters at 30% linkage distance. Cluster analysis was performed by Tewodros et al.
(2014) to study the patterns of groupings of fourteen bread wheat genotypes and grouped into
three clusters.

Cluster analysis based on agro-morphological traits of modern varieties and land races revealed
that the local varieties (land races) had the highest thousand-seed weight and unique phenotypic
characteristics in terms of ear shape and awn conditions as compared to other bread wheat
varieties (Mideksa et al., 2014). Awale and Sentayehu (2013) grouped twenty-six bread wheat
genotypes into six clusters using D2 analysis.

21
 3. MATERIALS AND METHODS

3.1. Description of the Study Area

The field experiment was conducted at Sirinka Agricultural Research Center (SARC) testing
sites of Jamma and Geregera

1. Jamma which lies between the geographical coordinates 10o 23’ to 10o 27’ N latitudes and 390
7’ to 390 24’ E longitudes in South Wollo Zone of the Amhara National Regional State, which is
260 km away from the capital city, Addis Ababa, in the north east direction and at geographical
coordinates of 100 27’ N latitude and 390 15’ E longitudes at an altitude of 2600 m.a.s.l. The
dominant soil type is of PH 6.0 with total rainfall of 720.5 mm, and minimum and maximum
temperatures of 10.0 and 21.1 0C, respectively.

2. Geregera is found 651 km away from Addis Abeba and located at an altitude of 2650-2855
m.a.s.l, which lies between 39 o N longitude and 12o E latitude with annual rainfall of 1105 mm.
The soil type is characterized as Lithosol, brown colour and pH of 5.6. Rainfall is erratic in
distribution, often unpredictable and is uni-modal, which starts in the first week of July and stops
at the end of August.

3.2. Planting Materials

Forty-nine bread wheat genotypes which are released varieties and elite materials taken from
Sinana and Kulumsa Agricultural Research Centers were used in the study. Description of the
genotypes are presented in Table 2.

22
 Table 2. Description of bread wheat genotypes used in the study

SN Genotype Code Pedigree Source Year of

center release
1 Honqolo - - Kulumsa -
2 Biqa - - Kulumsa -
3 WORRAKATTA/PAS - - Sinana 2014
TOR
4 UTQUE96/3/PYN Sinana 2014
/BAU//MILLAN
5 Hidasse ETBW5795 YANAC/3/PRL/SARA//TSI/VEE#5/4 Kulumsa 2012
6 Ogolcho ETBW 5520 WORRAKATTA/2*PASTOR Kulumsa 2012
7 Hoggana ETBW 5780 PYN/BAU//MILAN Kulumsa 2011
8 Hulluka ETBW5496 Kulumsa 2012
9 Mekelle-3 M17SAWSN79 - Mekele 2012
10 Mekelle-4 - - Mekele 2013
11 Shorima ETBW 5483 UTQUE96/3/PYN/BAU//MILAN Kulumsa 2011
12 Kakaba Picaflor#1 KIRITATI//SERI/RAYON Kulumsa 2010
13 Danda'a Danphe#1 KIRITATI//2*PBW65/2*SERI.1B Kulumsa 2010
14 Gassay HAR 3730 PASTOR Adet 2007
15 Alidoro HK14R251 HK-14-R251 Holeta 2007
16 Digelu HAR3116 SHA7/KAUZ Kulumsa 2005
17 Tay ET12/604 ET12D4/4777(2)//FKN/GB/3/PVN"S" Adet 2005
18 Sofumar HAR 1889 LIRA 'S'/TAN"S" Sinana 1999
19 Mada-Wolabu HAR 1480 TI/3/Fn/Th/Nar 59 *2/4/Bol'S' Sinana 1999
20 Pavon-76 - VCM//CNO"S"/7C/3/KAL/BB Kulumsa 1982
21 Jeferson - - Kulumsa 2012
22 King Bird - (300/SM+501M)/HAR 1709 Kulumsa 2014
23 ETBW 6861 - WAXWING*2/HEILO Kulumsa Pipe line
24 ETBW 8506 - AGUILAL/FLAG-3 Kulumsa Pipe line
25 ETBW 8507 - DURRA-4 Kulumsa Pipe line

23
26 ETBW 7120 - QAFZAH-23/SOMAMA-3 Kulumsa Pipe line
27 ETBW 8508 - REYNA-8 Kulumsa Pipe line
28 ETBW 7213 - CHAM4/SHUHA'S'/6/2*SAKER/5/R Kulumsa Pipe line
BS/ANZA/3/KVZ/HYS//YMH/TOB
29 ETBW 8509 - REYNA-29 Kulumsa Pipe line
30 ETBW 7038 - ATTILA/3*BCN//BAV92/3/TILHI/5/ Kulumsa Pipe line
BAV92/3/PRL/SARA//TSI/VEE#5/4/
CROC_1/AE.SQUARROSA
(224)//2*OPATA
31 ETBW 8510 - HIJLEEJ-1 Kulumsa Pipe line
32 ETBW 7058 - ROLF07//TAM200/TUI/6/WBLL1/4/ Kulumsa Pipe line
HD2281/TRAP#1/3/KAUZ*2/TRAP//
KAUZ/5/TACUPETO F2001
33 ETBW 8511 - BOW#1/FENGKANG Kulumsa Pipe line
15/3/HYS//DRC*2/7C
34 ETBW 7147 - CROC-1/AE.SQUARROSA(224)// Kulumsa Pipe line
OPATA/3/QAFZAH21/4/SOMAMA-
3
35 ETBW 8512 - BABAX/LR42//BABAX*2/3/KURU Kulumsa Pipe line
KU/4/KINGBIRD #1
36 ETBW 7871 - PAURAQ/4/PFAU/SERI.1B//AMAD/ Kulumsa Pipe line
3/WAXWING
37 ETBW 8513 - MUTUS//WBLL1*2/BRAMBLING/3 Kulumsa Pipe line
/WBLL1*2/BRAMBLING
38 ETBW 6940 - UTIQUE 96/FLAG-1 Kulumsa Pipe line
39 ETBW 8514 - TUKURU//BAV92/RAYON/3/WBLL Kulumsa Pipe line
1*2/BRAMBLING/4/
40 ETBW 7368 - D. 56455 Kulumsa Pipe line
41 ETBW 8515 - BECARD/3/PASTOR//MUNIA/ALT Kulumsa Pipe line
AR84
42 ETBW 7364 - ACSAD1115 Kulumsa Pipe line

24
43 ETBW 8516 - KACHU/KIRITATI Kulumsa Pipe line
44 ETBW 7194 - VAN'S'/3/CNDR'S'/ANA//CNDR'S'/ Kulumsa Pipe line
MUS'S'/4/TEVEE-5
45 ETBW 8517 - FRNCLN*2/TECUE #1 Kulumsa Pipe line
46 ETBW 7101 - KAMB2/PANDION Kulumsa Pipe line
47 ETBW 8518 - SUP152/AKURI//SUP152 Kulumsa Pipe line
48 ETBW 7872 - QUAIU/5/FRET2*2/4/SNI/TRAP#1/3 Kulumsa Pipe line
/KAUZ*2/TRAP//KAUZ
49 ETBW 8519 - ATTILA/3*BCN*2//BAV92/3/KIRIT Kulumsa Pipe line
ATI/WBLL1/4/DANPHE
Source: Kulumsa Agricultural Research Center, 2015.

3.3. Experimental Design, and Trial Management

The experiment was laid out in 7x7 simple lattice design with two replications. The dimension of
an individual plot area was 1.2m width x 2.5 m length (3m2) with six rows for each entry. The
spacing between blocks, plots and rows were 1.5m, 0.4m and 0.2m, respectively. The
experimental field was well tilled and planting rows were prepared using hand pulled row-
marker. Planting was done with the seed rate of 150 kg/ha (45 g/plot). Diammoniumphosphate
(DAP) and Urea fertilizers were applied at the rate of 100 kg/ha. Urea in splits: first split (1/3)
and the second split (2/3) of the total dose at planting and mid tiller stages, respectively. All the
cultural and agronomic practices were applied as recommended and kept constant. Weeds were
removed manually as and when required. The data for characters studied were collected from the
four central rows for each plot.

3.4. Data collected

For quantitative characters, AGROVOC descriptors: Cereal crops, plant developmental stages,
plant physiology, agronomic characters, yield components, measurement, sampling was adopted
and data were recorded on plant and plot basis as described below.

25
3.4.1. Plant basis

Ten plants were randomly taken from the four central rows for recording the following
observations and the mean values for the treatment were computed:

1. Plant height (PH): The distance in cm between the ground level to the tip of the terminal
spikelet of ten plants (excluding the awns) at maturity.

2. Number of productive tillers per plant (NPP): The actual count of the fertile numbers of
tillers of ten plants (spike bearing) per plant.

3. Spike length (SL): Length measured in cm from base of spike to the tip of the highest spikelet
of ten plants (excluding the awns) in cm at maturity.

4. Number of spikelets per spike (NSPS): Total numbers of spikelet’s on main spike of all ten
plants from four rows were counted at the time of maturity and average was recorded.

5. Number of grains per spike (NGS): The actual count of the number of kernel per spike of all
ten plants after threshed manually at the time of harvest.

3.4.2. Plot basis

The data on the following attributing traits were collected on the basis of the central four rows
(2m2) in each plot.

1. Days to heading (DH): The number of days from date of sowing to the stage where 50% of
the spikes have fully emerged.

2. Days to maturity (DM): The number of days from sowing to the stage when 75% of the
plants in a plot have reached physiological maturity.

3. Grain filling period (GFP): The number of days from heading to maturity, i.e. the number of
days to maturity minus the number of days to heading.

4. Thousand Seed weight (TSW): Weight of 1000 seeds randomly taken from each plot in
gram.

26
5. Biological yield (BY) (kg/m2): The representative plants within the four central rows or from
2m2/ plot were harvested and weighed in kilograms at maturity, after drying at 70°C for 24 hours
in a well-ventilated oven.

6. Grain yield (GY) (qt/ha): Grain yield in grams (g/m2 ) obtained from the central four rows
(2m2) of each plot and converted to quintals per hectare after moisture of the seed is adjusted
to 12.5% moisture content.

7. Harvest index (%): The ratio of yield of dried grain weight to the dried above ground
biomass weight of the harvestable plot (2m2 /plot) multiplied by 100.

3.5. Statistical Analysis

3.5.1. Analysis of Variance (ANOVA)

Analysis of variance was applied in order to test the significance differences of traits. The data
collected for each quantitative trait were subjected to analysis of variance (ANOVA) for simple
lattice design using Proc lattice and Proc GLM procedures of SAS version 9.2, (SAS Institute,
2008). Then after testing the ANOVA assumptions, Fisher’s protected least significant difference
(LSD) test at 5% level of significance was used for genotypes mean comparisons, whenever
genotype differences were significant. To perform a combined statistical analysis across
locations, test of homogeneity of error variances of each character for the two locations were
performed by using F- test (the ratio of the largest to the smallest error variance) to the
characters, and the test showed homogeneity of the two locations for all characters that involved
in the study. The ANOVA was also run for the two locations separately and combined over the
two locations since all characters showed homogeneity of error variance. The difference between
treatment means was compared using Fisher’s protected least significant difference (LSD) test at
5% probability levels. GENRES Version 7.01, (Pascal Institute, 1994) was employed for
estimation of correlation between traits, and path coefficient analysis.

27
The model for lattice design is:

Yil(j)  µ  ti  rj  (b | r)l(j)  eil(j)

Where; Yil(j) is the observation of the treatment i (i = 1, ..., v = k2), in the block l (l = 1, ..., k) of
the replication j (j = 1, ..., m);

µ is a constant common to all observations;

ti is the effect of the treatment i;
rj is the effect of the replication j;
(b|r)l(j) is the effect of the block l of the replication j;
eil(j) is the error associated to the observation Yil(j), where eil(j) ~ N(0, s), independent.

Table 3. Skeleton for individual location analysis of variance for simple lattice design

Source of variance Df Sum of squares Expected Mean F-Values

Squares

Replication (r) r-1 SSR MSr MSR/ MSe

Genotypes (g) - [Un adj.] g²-1 SSg MSg MSg/ MSe

- [adj.] g²-1 SSg MSg MSg/ MSe

Block within replication (b) r(b-1) SSB MSb MSb/MSe
[adj.]
Intra-block error (Ebe) (b-1) (rb-b-1) SSE MSe
Total rb2-1 SSt

28
Table 4. Analysis of variance in the case of a series of genotypes evaluated across
environments

Source of variation Degrees of Mean Expected Mean Squares

freedom Squares

Environment e-1 MSE σ2e +g σ2r(E) + r σ2GE + rg σ2E

Rep(block) (r-1)e MSR σ2e +g σ2r(E)

Genotype g-1 MSG σ2e + r σ2GE+ re σ2G

Genotype × (g-1)(e-1) MSGE σ2e + r σ2GE

Environment

Error (g-1)(e-1)e MSE σ2 e

Notes; Total phenotypic variance: Var (Yijk) = σ2p + σ2e + σ2GE+ σ2G

Phenotypic variance of genotypic means: Var (Yijk) = σ2p = σ2e + σ2GE + σ2G = MSG
re e re
Genotypic variance = σ G = (MSG -MSGE)/re
2

Heritability on individual experimental unit basis: H2 = σ2G

σ2G + σ2GE + σ2e
Heritability on a genotypic-mean basis: H2 =σ2G
σ2G + σ2GE + σ2e
e re
The above calculations were done according to Comstock and Robinson, (1952).
Source; FAO, (2009).
where: r = number of replication, G = number of genotypes, df = degree of freedom, b
= block, e= environment, Ebe= Intra-block error, SS = Sum of squares, MS = mean
squares, SSR and MSR are sums of squares and mean squares of replication,
respectively; SSG and MSG are sums of squares and mean squares of genotypes,
respectively; SSb and MSb are sums of squares and mean squares of blocks within
replication, respectively; SSe and MSe are sums of squares and mean squres of intra-
block error, respectively; and SSt is sum of squares of the total; σ2g=variance due to
genotypes, σ2 Ge= variance due to genotypes with environment interaction,
σ2e=variance due to environments.

29
3.5.2. Genotypic and Phenotypic Coefficients of Variation

Phenotypic and genotypic coefficients of variation were estimated according to Singh and
Chaudhary (1985).

 2g  2P
GCV  100 PCV   100
X X

Where; GCV = Genotypic coefficient of variation;

PCV =Phenotypic coefficient of variation
x̄= Grand mean of the characters under study

The mean values were used for genetic analyses to determine genotypic coefficient of variation
(GCV) and phenotypic coefficient of variation (PCV) (Singh and Chaudhury, 1985).

3.5.3. Genetic Advance (GA)

Genetic advance (GA) was calculated with the method suggested by Allard (1960); Singh and
Chaudhury (1985):
GA=k. σ2p. H2

Where; GA: genetic advance.

K: constant = 2.06 at 5% selection intensity.
σ2p: square root of phenotypic variance.
H2: Heritability in broad sense

3.5.3.2. Genetic advance as percent of mean

Genetic advance as percent of the mean was calculated to compare the extent of predicted
advance of different traits under selection, using the following formula:
GAM = GA x100
x̄
Where; GAM=Genetic advance as percent of mean
GA=Genetic advance under selection, and x̄=Grand mean of the trait

30
3.5.4. Correlation Analysis

Estimation of correlation coefficients (r) was computed using GENRES statistical software
(Pascal Intl Software Solutions, 1994). Phenotypic and genotypic correlations were estimated
using the standard procedure suggested by Miller et al. (1958) and Kashiani and Saleh (2010)
from the corresponding variance and covariance components.
gcovx.y pcov x.y
rg  rp 
 gx. gy
2 2
 2px. 2py

Where: rg = Genotypic correlation coefficient

rp = Phenotypic correlation coefficient

Gcovxy = Genotypic covariance between variables x and y

Pcovxy = Phenotypic covariance between variables x and y

2
σ gx = Genotypic variance for variables x
2
σ gy = Genotypic variance for variables y
2
σ px = Phenotypic variance for variables x
2
σ py= Phenotypic variance for variables y
To test the significance of correlation coefficients, the following formula was used (Sharma,
1998):

t=r/ SE(r)
Where; SE(r) =1-r2
Where; r is correlation coefficient
n is number of genotypes.
Then, calculated ‘t’ value was compared with standard value at n-2 degrees of freedom and a
levels of probability (where t is 0.05 and 0.01).

3.5.5. Path Coefficient Analysis

The analysis was done following the method suggested by Dewey and Lu (1959):

Rij= pij +∑ rik +pkj

31
Where rij = mutual association between the dependent character, i (yield-related trait) and
independent character, j (grain yield) as measured by the correlation coefficients; Pij is the
components of direct effects of the independent character (i), Σ rik pkj = summation of
components of indirect effect of a given independent character (i) on the given dependent
character (j) via all other independent characters (k). Whereas the contribution of the remaining
unknown characters is measured as the residual effect (R2) which is calculated as:
√ (1-R2)
Where, R2=∑ pij +rij

3.6. Principal Component Analysis

Principal component analysis was performed using correlation matrix by employing PAST 1.93
(Palaeontological Statistics; Hammer et al., 2001) to evaluate the contribution of each
quantitative character in the total variation of genotypes. Number of factors retained was decided
by looking at the Eigen values (values > 1.0) (Fellahi et al., 2013). Those traits that had load
coefficient values > 0.40 (ignoring the sign) were considered as relevant scores for the PCAs.
The general formula to compute scores on the first component extracted (created) in a principal
component analysis is described as:

C1 b11 X 1  b12  b1 p Xp 

Where, C1 = the subject’s score on principal component 1 (the first component extracted)
b1p = the regression coefficient (or weight) for observed variable p, as used in creating
principal component one.
Xp = the subject’s score on observed variable p.

3.7. Genetic Divergence and Cluster Analysis

Mahalanobis (1936) statistics was used to estimate the genotypic divergence between clusters.
All the genotypes used were clustered into different groups based on D2 statistics. The D2 values
of all the combinations were arranged in descending order. D2 statistics is defined by the
following formula:

D2ij= (Xi -Xj) S-1 (Xi – Xj)

Where; D2ij = the square distance between any two genotypes i and j;

32
 Xi and Xj = the vectors for the values for genotypes ith and jth genotypes;
S-1 = the inverse of pooled variance covariance matrix within groups.
Testing the significance of the squared distance values obtained for a pair of clusters was taken
as the calculated value of c2 (chi-square) and tested against the tabulated c2 values at p-1 degree
of freedom at 1% and 5% probability levels, where p = number of traits used for clustering
genotypes.

The proc cluster of SAS system with average linkage method of clustering strategy version 9.2
(SAS Institute, 2008), which grouped and sorted the genotypes into clusters to form Dendrogam.
Cubic clustering criterion (CCC), pseudo F (PSF), and pseudo t2 (PST2) statistics were used in
determining the number of clusters in the data.

33
 4. RESULTS AND DISCUSSION

4.1. Analysis of Variance

The analysis of variance for different characters at Jamma and Geregera locations are presented
in Appendix table 2, and 3 respectively. There was significant differences at (P< 0.01 and P<
0.05) levels among genotypes for all characters considered the two environments except for grain
filling period at Geregera, which was non-significant.

Since the relative efficiency of simple lattice design is less than complete randomized block
design (RCBD) for most characters (Appendix table 2 and 3), which showed under 105% and
also blocks within replication sum of squares were non-significant. Therefore analysis of
variance were performed using complete randomized block design (RCBD) model. Similar
findings were reported by Azeb et al. (2016). Before pooling the data across environments, test
of homogeneity using F-test for error of variance was done. Therefore, the hypothesis of
homogeneous variance is accepted, and analysis of variance and other statistical analysis were
run for combined over the two locations.

Combined analysis of variance results for different studied traits is shown in (Table 5). The
location effect was significant for all traits, indicating the different climatic conditions in two
locations. The location × genotype interaction effect was significant for all traits except number
of spikelets per spike indicating different performance of bread wheat genotypes across the two
locations. Furthermore, Mean square of genotypes for all characters studied were significant
(P<0.05 and P<0.01) differences among the bread wheat genotypes, indicating the existence of
sufficient genetic variability within different genotypes to be exploited in the breeding programs
that was also reflected in the broad ranges observed for each traits as presented in (Table 5),
representing the genetic diversity for further selection procedures in the experimental material
under study. The present investigation are in confirmation with early findings of (Ashamo et al.,
2012; Awale et al., 2013; Kumar et al., 2014; Tewodros et al., 2014; Zeeshan et al., 2014;
Adhiena et al., 2016).

34
 Table 5. Estimated values of mean squares C.V (%) and R-square (%) for 11 traits of
49 bread wheat genotypes combined over across locations

Traits Sources of Variance

E Re(b) G ExG error C.V (%) LSD at 5% R-Square (%)

DF 1 1 48 48 96

DH 65.15** 5.45ns 58.12** 10.5** 3.110 2.65 2.480 0.92

DM 650.3** 0.785ns 34.67** 11.70* 5.900 1.90 3.410 0.84

PHT 4662** 348.6** 117.4** 44.7** 25.50 6.45 7.090 0.84

NPTP 9.48** 1.77** 0.163** 0.10** 0.060 16.3 0.350 0.81

SL 99.26** 13.17** 2.40** 0.710* 0.460 8.97 0.940 0.86

NSPS 321.4** 9.130* 3.50** 1.30ns 1.130 7.30 1.480 0.84

NGS 922.4** 2.800ns 40.5** 17.00* 11.27 8.90 4.710 0.77

TSW 9839** 15.00ns 41.30** 16.15* 6.800 7.11 3.900 0.94

BY 23.40** 0.020ns 0.140** 0.088** 0.045 10.0 0.299 0.89

HI 1.070** 0.006* 0.00311** 0.0021* 0.0014 9.66 0.043 0.94

GY 35311.8** 75.60* 61.72** 43.75** 22.50 13.7 6.66 0.95

NB: E=environment, Re(b)= replication within a block, E x G= environment with Genotype

interaction mean square, CV=coefficient of Variation, DF=degrees of freedom, DH=Days
to heading, DM=Days to maturity, PHT=plant height, NPT=number of productive tillers
plant, SL =Spike length, NSPS=Number of spike lets per spike, NGS=Number of grains
per spike, BY=Biological yield, HI=Harvest index, TSW=Thousand seed weight, and
GY=Grain yield per hectare.

4.2. Genetic Variability and Mean Performance of Genotypes

The success of a breeding program depends largely upon the amount of genetic variability
present in the population and the extent to which the desired traits are heritable. Based on

35
combined over location the mean performance of genotypes for studied traits showed a wide
range of variation (Table 6). Days to heading ranged from (61-79.5 days ), with mean value of
66.6 days, days to maturity (124-136 days) with mean value of 127.6 days, plant height ranged
from 68-93.75 cm with mean value of 78.3 cm, productive tillers per plant(1.2-1.9) with mean
value of 1.5, spike length (6.4-10.9 cm) with mean value of 7.5 cm, number of spikelets per
spike(13-17.4) with mean value of 14.6, number of grains per spike (29.45-7) with mean value of
37.8, , thousand seed weight (34.8-48 g) with mean value of 40.8 g, biological yield (1.8-2.7 kg)
with mean value of 2.12 kg, harvest index (0.26-0.36) with mean value of 0.31, grain yield per
hectare (26.5-43.8 qt/ha) with mean value of 34.6 qt/ha, indicating good opportunity for grain
yield improvement. A wide range of variation among bread wheat genotypes in yield and yield
related traits reported (Ashamo et al., 2012; Awale et al., 2013; Kumar et al., 2014; Tewodros et
al., 2014; Adhiena et al., 2016).

Based on mean performance of genotypes the response of grain yield for separate and across
location was discussed below. At Jamma environment, grain yield ranged from Genotypes such
as ETBW 8518 (60), Mada-wollabo (58.5), ETBW 8506 (57), Hoggana (56.25), Ogolcho (55)
were better grain yielder in Qt/ ha respectively. Whereas genotypes such as Gassay (32.5),
ETBW 7058 (32), Biqa (29.25), Mekele-3 (27.5) and Mada-wollabo (27.5) were better grain
yielder in Qt/ ha under Geregera respectively. Genotypes such as Gassay (44.75), Mada-
wollabo (43.00), Biqa (41.5), Mekele-3 (40), UTQUE96/3/PYN/BAU//MILLAN (39.9) were the
top grain yielder genotypes across location. Thus it was observed that the overall mean for grain
yield was the lowest (21.17 qt/ha) at Geregera environment, whereas Jamma seems to be ideal
for cultivation of bread wheat as the overall mean grain yield of the location was (48 qt/ha) as
data presented in appendix (Table 2 and 3).

4.3. Genotypic and Phenotypic Coefficients of Variation

Because of high genotype-environment (G × E) interactions, estimates of GCV, H2 and GA for

most of the characteristics using combined over location analysis were generally lower than the
estimates computed from the variance analyses made separately for each location as presented in
(Table 6 and Appendix table 2, 3) respectively.

36
Low genotypic as well as phenotypic coefficient of variation in the characters observed may be
due to presence of both positive and negative alleles in the population (Majumder et al., 2008).
The genotypic coefficient of variation ranged from 1.88 % for maturity date to 8.66 % for spike
length; and phenotypic coefficient of variation ranged from 2.3% for maturity date to 13.3% for
number of productive tillers per plot (Table 6). Maximum values of genotypic coefficient of
variation were recorded for spike length (8.66 %), followed by number of productive tillers per
plant (8.4%), number of grains per spike (6.4 %), and thousand seed weight (6.15 %), whereas
better value of phenotypic coefficient of variation were recorded for productive tillers followed
by grain yield, spike length, and harvest index with a value of 13.3%, 11.35 %, 10.3%, 9%
respectively in the study.

The magnitude of phenotypic coefficient of variation (PCV) is much higher than the genotypic
coefficient of variation (GCV) for number of productive tillers per plant, grain yield, harvest
index and biological yield indicating that apparent variation for the characters was not only due
to genotypes but also due to influence of wide range of phenotypic (VP or σ2P) and genotypic
variance (VG or σ2g) observed in the experimental material for all the traits studied. This result is
related with the findings of other similar works (Kashif et al., 2004; Subhani et al., 2010;
Mohammedi et al., 2011; Asaye et al., 2013). Likewise, the phenotypic variances for plant height
and days to heading were also high, indicating that the genotype could be reflected by the
phenotype and the effectiveness of selection based on the phenotypic performance for these
traits.

4.4. Estimation of Heritability in Broad Sense

The concept of heritability explains whether differences observed among individuals arose as a
result of differences in genetic makeup or due to environmental forces (Azeb et al., 2016). More
effective in breeding of homozygous lines. Heritability estimate for characters under study is
indicated in (table 6). In this study heritability in broad sense ranged from 29% for grain yield to
82% for heading date. The heritability is categorized as low (0-30%), moderate (30-60%) and
high (60% and above) as given by Comstock and Robinson, (1949). Accordingly, high
heritability was estimated for days to heading (82%), maturity date (66.2%), spike length
(70.4%), plant height (63.6), number of spikelets per spike (62.5) and thousand-seed weight
(61%). Similar results were reported by Laghari et al. (2010); and also Ali et al. (2008) and

37
Karim et al. (2013) reported high estimates of heritability for spike length and number of
spikelets per spike in bred wheat.

Moderate heritability was obtained for number of grains per spike, number of productive tillers,
harvest index and biological yield, indicating that the characters were more influenced by
environment. Although high heritability estimate have been found to be effective in the selection
of superior genotypes on the basis of phenotypic performance, Kumar et al., (2014), suggested
that heritability estimates along with genetic advance will be more useful in predicting the effect
for selecting the best individual.

Related findings were reported by Laghari et al. (2010). Low heritability was obtained for yield
per hectare (29%) which is in agreement with the results obtained by (Mollasadeghi et al., 2012;
Mohammadi et al., 2011; Tesfaye et al., 2014; Mesele et al., 2016) with a value of (12.27%,
7.4%, 19% and 25%) respectively. Opposed to this study, Awale et al. (2013) reported low
estimates of heritability for spike length, while Tewodros et al., (2014) reported low estimates of
heritability for heading date (13%), maturity date (7.79%), plant height (12.8%) and thousand-
seed weight (32.8%). The main difference in the findings may be due to the difference in the
genetic material used and environmental conditions (Kashif et al., 2004; Mohammadi et al.,
2011; Adhiena et al., 2016).

Heritability determine the choice of plant breeding method/technique. High h2 use mass and pure
line selection whereas low h2 use recurrent selection. For traits with low h2 selection in early
segregating generation (F2) would be effective because of in subsequent generation’s variation
decreases due to increase of homozygosis. High heritability accompanied with relatively high
genetic advance in case of, plant height, spike length, and thousands seed weight indicates that
most likely the heritability is due to additive gene effects and selection may be effective in early
generations for these traits. High heritability for days to heading, days to maturity, number of
spikelets per spike coupled with low genetic advance indicates non-additive gene effects.
Therefore, there seems a limited scope for improvement in this trait.

38
4.5. Estimates of Expected Genetic Advance (GA)

Genetic advance as a percent mean ranged from 3.15% for maturity date to 14.9% for spike
length (Table 6). Relatively high genetic advance as a percent mean were recorded for spike
length followed by number of productive tillers per plant, number of grains per spike, thousand
seed weight, heading date and plant height with values of 14.9%, 10.6%, 10%, 10%, 9.7%, and
9.07%respectively, indicating good response to selection.The present study was in close
agreement with the findings of (Mohammadi et al., 2011; Asaye et al., 2013; Adhiena et al,
2016; Rahman et al., 2016).

It was suggested that the importance of considering both the genetic advance and heritability of
traits rather than considering separately in determine how much can progress to be made via
selection (Kumar et al., 2014). As a result, traits like spike length, number of grains per spike,
and thousand seed weight showed high heritability accompanied with better genetic advance as
percent of mean and genetic and phenotypic coefficient of variation in this study. The high
heritability estimates along with low genetic advance indicates that non additive type of gene
action and genotype-environment interaction plays a significant role in the expression of the
traits as observed in days to maturity in the present study, which agrees with the findings of
(Majumder et al., 2008).

Plant height, spike length, grains per spike and thousand seed weight weight had relatively high
heritability along with high genetic advance in percentage of mean making them most important
in the selection of modern wheat. High GCV, PCV, heritability and GA% of mean for spike
length suggested that it could be transmitted to the hybrid progeny and phenotypic selection
based on this would be effective.

39
 Table 6. Estimates of range, means, genotypic and phenotypic v ariances, broad sense
heritability, genetic advance, and genetic advance as a percentage of mean for 11
characteristics of 49 bread wheat genotypes, combined across the locations

Traits Range Mean ± SE δ2g δ2 p GCV (%) PCV (%) H2 (%) GA GAM
DH 61-79.5 66.6±0.04 11.90 14.53 5.20 5.72 82.0 6.45 9.70
DM 124-136 127.6±0.22 5.74 8.67 1.88 2.30 66.2 4.02 3.15
PHT 68-93.75 78.3±0.084 18.7 29.35 5.50 6.90 63.6 7.10 9.07
NPT 1.2-1.95 1.5±0.214 0.016 0.040 8.40 13.3 38.7 0.16 10.6
SL 6.4-10.9 7.5±0.104 0.422 0.600 8.66 10.3 70.4 1.12 14.9
NSPS 13 -17.4 14.6±0.78 0.550 0.880 5.10 6.44 63.0 1.21 8.30
NGS 29-45.7 37.8±0.11 5.850 10.12 6.40 8.42 57.8 3.80 10.0
TSW 34.8-48 40.8±0.10 6.29 10.32 6.15 7.86 61.0 4.04 10.0
BY 1.8-2.70 2.12±0.123 0.013 0.035 5.38 8.82 37.0 0.143 6.73
HI 0.26-0.36 0.31±0.16 0.00025 0.00078 5.10 9.00 32.0 0.018 5.95
GYP 26.5-43.8 34.6±0.20 4.50 15.43 6.13 11.35 29.1 2.360 6.80
NB: σ2g = genotypic variance, 2
σ p=phenotypic variance, GCV (%) = genotypic coefficient of
variation, PCV (%) = phenotypic coefficient of variation, H2 (%) =broad sense heritability,
GA =genetic advance, GAM % =genetic advance as percentage of mean, DF=degrees of
freedom, DH=days heading, DM=days maturity, PH=plant height, NPTP=number of
productive tillers per plant, SL =spike length, NSPS=number of spikelets per spike,
NGS=Number of grains per spike, TSW= thousand seed weight, BY=biological yield,
HI=harvest index, and GY= Grain yield.

4.6. Correlations Analysis of Quantitative Traits

Genotypic and phenotypic correlations of all possible combinations for traits under study are
presented in (Table 7), provided that in most of the cases the genotypic correlation coefficient
were higher than the corresponding phenotypic correlation coefficient indicating strong inherent
relation between the traits but suppressing effect of the environment, which modified the
phenotypic expression of these characters by reducing phenotypic coefficient values.

40
A positive value of r (correlation) shows that the changes of two variables are in the same
direction, that is, high values of one variable are associated with high values of other and vice
versa (El-Mohsen et al., 2012). In general the magnitude of genotypic correlations (rg) is higher
than those of phenotypic correlations (rp). This revealed that association among characters is
under genetic control and indicating the preponderance of genetic variance in expression of
characters. It might be due to depressing effect of environment on character association as
reported earlier for wheat crop (Laghari et al., 2010; El-Mohsen et al., 2012). When value of rp is
greater than rg, it shows apparent association of two traits is not only due to genes but also due to
favorable influence of environment. By contrast, if value of r is zero or insignificant, this shows
that the two traits are independent.

Thus from the study, positively and significantly correlation of characters studied with grain
yield per hectare both at genotypic and phenotypic levels, suggests that yield per hectare would
increase with increase of those characters and vice versa.

Days to heading: Days to heading showed negative non-significant association at genotypic and
at phenotypic levels (rg = -0.184, rp = -0.102) with grain yield per hectare. El-Mohsen et al.
(2012) and Awale et al. (2013) reported negative associations between days to heading and grain
yield per plot at genotypic and phenotypic levels. While Moghaddam et al. (1997) and Ali et al.
(2009) reported positive association between days to heading and grain yield per plot. Days to
heading highly significant positive association at genotypic and at phenotypic levels with
maturity date and highly negative associated with number of productive tillers per plant at
genotypic level.

Days to Maturity: Days to maturity showed negative association at genotypic levels (rg = -
0.252) and at phenotypic level (rp =-0.021) with grain yield per hectare. This finding is in
agreement with the findings of Awale et al. (2013) and contradicted with the findings of Ali et
al. (2009). The findings of Khan et al. (2010) showed positive association at genotypic levels
and negative association at phenotypic levels. Days to maturity negative significant associated
with biological yield per plot at genotypic level. On the other hand it was positive non-significant
associated with spike length, number of spike lets per spike, and thousand seed weight at both

41
levels. While negatively non-significant associated with other traits at both levels, except harvest
index at phenotypic level.

Plant height: The correlation between plant height and grain yield per hectare was positive and
significant at both genotypic and phenotypic levels (rg = 0.384**, rp = 0.354*) which indicates
that an increase in plant height leads to an increase grain yield. Similar results have been found
(Moghaddam et al., 1997; Kashif and Khaliq, 2004; Aydin et al., 2010; Fellahi et al., 2013;
Farshadfar et al., 2014; Awale et al., 2013; Gelalcha and Hanchinal, 2013). However, El-Mohsen
et al. (2012) reported negative correlation of plant height and grain yield. Plant height showed
negative non-significant association at genotypic and at phenotypic levels with days to heading
and days to maturity. However it was positive non-significant associated with number of
productive tillers per plant and harvest index at both correlation types. Moreover it was highly
associated with spike length, number of spikelets per spike, number of grains per spike, thousand
seed weight and biological yield.

Number of productive tillers per plant: The correlation between number of tillers per plant
and grain yield per hectare was positive and significant at both genotypic and phenotypic levels
(rg = 0.366*, rp = 0.226). Number of tillers per plant was negatively and highly significant
associated with number of spikes per spike at genotypic level (rg = -0.381**), and also non-
significant negative correlation with spike length, suggesting that increase in tiller number
reduce , number of spikes per spike and spike length, which are similar with El-Mohsen et al.
(2012). Number of productive tillers per plant displayed positive and significant relationship at
genotypic level with thousand seed weight, and positive and non-significant relationship at
genotypic level with number of grains per spike, biological yield and harvest index, suggesting
that increase in tiller number adds the value of those traits, indicated that number of tillers per
plant may be an effective trait to select higher yielding genotypes.

Spike length: Spike length was in negative relationship at genotypic levels and in positive
phenotypic with grain yield per hectare (rg = -0.047, rp = 0.014). These results are supported by
the findings of earlier researchers like Khan et al. (2010). A positive and highly significant
correlation was observed between spike length and number of spikelets per spike. It means that
with the increase in spike length there was a significant increase in number of spikelets per spike

42
as discussed by Ul-haq et al. (2010). There was a positive correlation between spike length and
thousand seed weight at genetic level and also positive and highly significant correlation was
observed between spike length and plant height.

Number of spikelets per spike: Number of spikelets per spike was in positive relationship at
genotypic level (rg = 0.004) and in negative relationship at phenotypic level (rp = -0.124) with
grain yield per hectare. A significant and positive phenotypic correlation was observed between
numbers of spikelets per spike and plant height, number of productive tillers per plant and at
genotypic level highly correlated with numbers of grains per spike. Kashif and Khaliq (2004)
and El-Mohsen et al. (2012) also observed number of spikelets per spike as significantly and
positively correlated with grain yield at genotypic level.

The number of spikelets per spike showed negative and highly significant correlation with spike
length and number of grains per spike at the genotypic level which agrees with the findings of
(Awale et al., 2013, Ali et al., 2009), while positive highly significant correlation with spike
length at the phenotypic level also agrees with (Ali et al., 2009).

Number of grains per spike: It had positive association with grain yield per hectare at
genotypic level (rg = 0.176**), and at phenotypic level (rp = 0.136). It had highly significant
positive relationship with plant height at genotypic and phenotypic level. The perusal of both the
correlation coefficient results suggested that number of grains per spike should be given prime
importance regarding its contribution to yield. These results suggest that selections should be
based on number of grains per spike for developing new high yielding wheat varieties. These
results are substantiated with those of Kashif and Khaliq (2004) and El-Mohsen et al. (2012).

Thousand seed weight: Thousand seed weight showed positive and significant association at
genotypic and phenotypic levels (rg = 0.395*, rp = 0.365) with grain yield per hectare. This result
is in agreement with a number of works in wheat (Kashif et al., 2004; Khaliq et al., 2004;
Mohibullah et al., 2011; Iftikhar et al., 2012; Kalimullah et al.,2012; Laei et al., 2012;
Zafarnaderi et al., 2013), but contradicted with the findings of Khan et al.(2010) and Awale et
al. (2013). The interrelation between yield contributing characters exhibits that thousand seed
weight was positively correlated with harvest index which indicated high portion of
photosynthesis was due to increase thousand seed weight.

43
Biological yield: It was in positive and highly significant relationship at both phenotypic and
genotypic levels with grain yield per hectare (rg = 0.617**, rp = 0.624**). These results are
supported by the findings of Chowdhry et al. (1991), Laei et al. (2012) and Chimber et al.
(2014). Also, it was highly and positively correlated with plant height at both genotypic and
phenotypic levels. The results corroborate the findings of Moghaddam et al. (1997).

Harvest Index: Harvest index had positive and significant relationship at both genotypic and
phenotypic levels with grain yield per hectare (rg = 0.731**, rp = 0.625*). These results are
supported by the findings of Chowdhry et al. (1991), Laei et al. (2012) and Zafarnaderi et al.
(2013). It was negatively correlated with days to heading, days to maturity, spike length and
thousand seed weight at genotypic level, and the result is supported by the findings of
Moghaddam et al., (1997), but contradicted with the findings of Zafarnaderi et al., (2013).

The significant correlation suggests that these traits could be used as indirect selection traits for
grain yield, i.e., increase of these traits would increase grain yield per hectare (Asaye et al.,
2013). The study of correlation among yield and yield contributing traits suggests that plant
height, number of productive tillers per plant thousand seed weight, harvest index and biological
yield were the most important characters which possessed highly positive association with grain
yield per plant. Therefore, these characters could be utilized in breeding program to improve
varieties for higher yield.

44
Table 7. Genotypic correlation coefficient (rg) (upper diagonal) and phenotypic correlation
coefficient (rp) (below diagonal) of 11 traits of 49 bread wheat genotypes

Traits DH DM PH NPTP SL NSPS NGS TSW BY HI GY

DH 0.946** -0.165 -0.386** 0.148 0.095 -0.092 0.207 -0.168 -0.122 -0.184
DM 0.767** -0.064 -0.099 0.155 0.107 -0.017 0.096 -0.306* -0.033 -0.252
PH -0.113 -0.039 0.26 0.565** 0.575** 0.625** 0.377** 0.363* 0.232 0.384**
NPTP -0.132 -0.033 0.18 -0.261 -0.381** 0.159 0.288* 0.268 0.248 0.366*
SL 0.087 0.107 0.474** -0.132 0.743** 0.032 0.218 -0.047 0.06 -0.047
NSPS 0.038 0.033 0.38** -0.039 0.662** 0.248 -0.012 0.046 -0.014 0.004
NGS -0.09 0.005 0.39** 0.064 0.007 0.188 0.019 0.061 0.177 0.176
TSW 0.154 0.161 0.372** 0.214 0.213 0.03 0.012 0.109 0.396** 0.395**
B -0.123 -0.057 0.375** 0.169 0.038 0.046 0.118 0.194 -0.067 0.617**
HI -0.031 0.031 0.161 0.15 0.021 -0.165 0.064 0.322* -0.144 0.731**
GY -0.102 -0.021 0.354* 0.226 -0.014 -0.124 0.136 0.365* 0.624** 0.625**
X2 =0.288, 0.372 (*, **) at 5 % and 1% probability level respective, DH=days to heading, DM=days to
maturity, PH=plant height, NPTP=number of productive tillers per plant, SL =spike length,
NSPS=number of spikelets per spike, NGS=Number of grains per spike, BYP=biological yield,
HI=harvest index, TSW= thousand seed weight, and GY= Grain yield.

45
4.7. Path Coefficient Analysis

Knowledge of correlation alone is often misleading as the correlation observed may not be
always true. Two characters may show correlation just because they are correlated with a
common third one. In such cases, it becomes necessary to use a method which takes into account
the causal relationship between the variables, in addition to the degree of such relationship. Path
coefficient analysis measures the direct influence of one variable upon the other, and permits
separation of correlation coefficients into components of direct and indirect effects. Partitioning
of total correlation into direct and indirect effects provide actual information on contribution of
characters and thus form the basis for selection to improve the yield.

Estimates of path coefficient analysis, direct and indirect effects of yield contributing characters
on grain yield per hectare using genotypic correlation, which showed significant association with
grain yield were presented in (Table 8). Maximum positive direct effect on grain yield per
hectare was exerted by harvest index (0.753), followed by biomass yield (0.753). The high direct
effects of these characters on grain yield could be considered as causes of such high correlation.
This means that a slight increase in one of these traits may directly contribute to grain yield.
Chowdhry et al. (1991) also reported positive direct effect of harvest index (0.443) and
biological yield (0.327) on grain yield per plant. On the other hand, negative direct effect was
exhibited by plant height (-0.215), number of productive tillers per plant (-0.078),

Plant height and number of productive tillers per plant showed negative direct effect on
grain yield by displaying a value of (-0.215, -0.078) respectively. Since the direct effect were
negative, so the direct selection for these trait to improve yield will be undesirable (Ali et al.,
2008), and positive indirect effect through biological yield per plot, thousand seed weight and
harvest index.

Thousand-seed weight vs. grain yield: Positive direct effect in case of thousand-seed weight on
grain yield was estimated by displaying a value of (0.161) on grain yield and in addition to this
thousand seed weight affected grain yield indirectly through harvest index (0.298*).

Biological yield and Harvest index showed positive direct effect on grain yield by displaying a
value of (0.753)

46
Dramatic increase in the grain yield of major world cereal crops is due mainly to increases in the
harvest index and to a lesser extent the biological yield (Acquaah, 2007). In this study both
harvest index and biological yield showed high genotypic correlation and positively significant
direct effect on grain yield. Thus plant breeder should practice selection through those most
favorable traits for future wheat yield improvement programs.

On the basis of estimates of path coefficients, it could be suggested that harvest index followed
by biological yield and thousand seed weight are the main contributors to grain yield in the
present investigation. The result agrees with Arega et al. (2007) and Gelalcha and Hanchinal
(2013), reported that traits such as biomass and harvest index, which showed highly significant
correlation with grain yield, can be used as selection indices in grain yield improvement.
Therefore, selection for characters will possibly improve other component characters thereby
improving grain yield.

The residual effect in path analysis determines how best the resultant component (independent)
variables account for the variability of the causal (dependent variable), grain yield per plant
(Singh and Chaudhary, 1985). To this end, residual effect in the present study was 0.126
showing that 87.4 % of the variability in grain yield was explained by the component factors.
This further indicates the interventions of environmental factors on the expression of the
characters for the choice of yield attributing traits. This result was related with the findings of
Gelalcha and Hanchinal (2013) and Arega et al. (2007), who reported residual effects 0.065 and
0.0083, respectively, indicating that characters included in the study explained high percentage
of variation in grain yield per plot. It also indicate that in addition to the studied characters, there
are also other factors to justify grain yield per plot changes (El-Mohsen et al., 2012).

47
Table 8. Estimate of direct (bold face and diagonal) and indirect (off diagonal) effects
at genotypic level in 10 traits of 49 bread wheat genotypes

Traits PH NPTP TSW BY HI rg

PH -0.215 -0.020 0.061 0.273 0.174 0.384**
NPTP -0.056 -0.078 0.046 0.202 0.187 0.366*
TSW -0.081 -0.023 0.161 0.082 0.298* 0.395**
BY -0.078 -0.021 0.018 0.753** -0.050 0.617**
HI -0.050 -0.019 0.064 -0.050 0.753** 0.731**
Residual effect= 0.126 *, ** indicate significance at 0.05 and 0.01 probability levels,
PH=plant height, NPTP=number of productive tillers per plant, TSW= thousand seed
weight, BY=biological yield, HI=harvest index, rg=genotypic correlation.

4.8. Principal Components Analysis

Principal component analysis (PCA) reflects the importance of the largest contributor to the total
variation at each axis of differentiation (Sharma, 1998). The main advantage of principal
component analysis is reducing the number of dimensions without much loss of information
(Fellahi et al., 2013). The eigenvalues are used to determine how many factors to retain and the
sum is usually equal to the number of variables (Daniel et al., 2011; Awale and Sentayehu, 2013
and Fellahi et al., 2013). The first step in PCA was to calculate eigenvalues, which all together
explained total variability that is displayed on the PC axes. The PCs with eigenvalue > 1.0 are
used as criteria to determine the number of PCs (Fellahi et al., 2013).

Data presented in Table 9 demonstrated that an increase in the number of components was
associated with a decrease in Eigen values. According to the results, the estimated wheat variable
had grouped into five principal components (PCs) such as PC1 (25.48%), PC2 (20.85%), PC3
(16.8%), PC4 (10.0), and PC5 (7.73) with Eigen values more than one (2.79, 2.18, 1.96, 1.15,
1.01) respectively, which all together explained 80.4% of total variability, leaving the remaining
19.4% in the last six principal components (Table 9).

Daniel et al., (2011) taken characters which load high positively or negatively with a value of
greater than ±3.0, contribute more to the variability and they are the ones that most differentiated

48
the clusters. Hence, data presented in Table 7 and graphically shown in Appendix figure 1
showed the most contributing characters are found in the first principal component which were
plant height, biomass yield, thousand seed weight and grain yield; whereas in the second PC
were days to heading, days to maturity, spike length and number of spikelets per spike; in the
third PC were harvest index, spikelets per spike, biological yield, grain yield; and in the fourth
PC were biological yield, number of grains per spike, harvest index and thousand seed weight
were the major contributing characters for variability to those principal components.

The factor loadings refer to the coefficients in each principle component or the correlation
between the component and the variables. A high correlation between PC1 and a variable
indicates that the variable is associated with the direction of the maximum amount of variation in
the data set. The components and their contributions in the variables are graphically shown in
Appendix figures 1. The present study confirmed the bread wheat genotypes showed wide
variations for the character studied and it suggests ample opportunities for genetic improvement
of bread wheat through direct selection from the genotypes, and conservation of the germplasm
for future utilization. Similar findings of grouping bread wheat genotypes by principal
component analysis were reported (Daniel et al., 2011; Awale and Sentayehu, 2013; Fellahi et
al., 2013).

49
Table 9. Vector loadings and percentage of explained variation by the first four PCs

Character PCA1 PCA2 PCA3 PCA 4 PCA 5

Hd -0.165 0.451 0.434 0.212 0.117

Md -0.112 0.432 0.456 0.158 0.223
PHT 0.483 0.204 -0.184 0.020 0.157
NPT 0.249 -0.259 0.141 0.057 0.221
SL 0.212 0.479 -0.245 -0.096 -0.332
NSPP 0.194 0.446 -0.364 -0.072 -0.081
NGS 0.255 0.055 -0.147 -0.096 0.811
TSW 0.319 0.122 0.357 -0.058 -0.201
BYP 0.337 -0.133 -0.023 0.7281 -0.126
HI 0.296 -0.082 0.364 -0.601 -0.097
GYP 0.461 -0.174 0.273 0.088 -0.153

Eigen value 2.800 2.260 1.800 1.100 1.010

% proportion 25.48 20.85 16.36 10.00 7.730

Cumulative% 25.48 45.22 63.04 73.49 80.40

NB: PCA=Principal component axis, DH=days to heading, DM=days to maturity, PH=plant

height, NPTP=number of productive tillers plant, SL =spike length, NSPS=number of
spikelets per spike, NGS=Number of grains per spike, BYP=biological yield, HI=harvest
index, TSW= thousand seed weight, and GY= Grain yield.

4.9. Genetic Divergence (Distance) Analysis

Divergence analysis is performed using Mahalanobis (1936) D2 distance to classify the diverse
genotypes for hybridization purpose (Table 10 and 11). The genetic improvement through
hybridization and selection depends on the extent of genetic diversity between parents. Chi-
square values were tested for significance using P-1 degrees of freedom where, P is the number
of characters used in the study (Singh and Chaudhary, 1985).

50
Inter cluster divergence values (D2) between and within seven clusters are presented in the (table
10). The highest inter-cluster distance was exhibited between cluster I and III (D2 = 25.79**),
followed by cluster II and IV (D2 = 22.82), and cluster II and III (D2 = 22.75), indicating wider
genetic divergence among the clusters. Thus, crossing of genotypes between members of cluster
I with members of cluster IV, and members of cluster II with members of cluster III, and IV may
produce a high amount of heterotic expression in the F1’s and broad spectrum of variability in
segregating (F2) populations. Genetic divergence in bread wheat genotypes reported by earlier
workers (Kashif et al., 2004; Ali et al., 2008; Daniel et al., 2011; Degewione and Alamerew,
2013; Fellahi et al., 2013).

Table 10. Inter and intra (bold) cluster D 2 values among six clusters in 49 bread wheat
genotypes

Cluster I II III IV V VI
I 10.29 25.79** 7.36 13.31 16.07
II 22.75* 22.82* 17.00 19.78*
III 8.910 12.22 15.68
IV 7.060 17.00
V 14.58
VI
X2 =18.3, 23.2 (*, **) at 5 % and 1% probability level respective

4.10. Clustering of Genotypes

The dendrogram obtained from the cluster analysis through average linkage technique grouped
the 49 genotypes into six clusters at about 47% similarity level based on D2 values considering
their pooled mean as data presented in (Table 11) and, as shown in (appendix figure 2) which
makes them moderately divergent. Related findings were reported by earlier workers (Daniel et
al., 2011; Awale and Sentayehu, 2013; Fellahi et al., 2013; Mideksa et al., 2014). Similarity
between clusters is the average distance between all objects in one cluster and all objects in other
cluster, where by individuals within any cluster were more closely related than individuals in
different clusters. The distribution of genotypes into different diversity classes of cluster
membership indicated that the genotypes are moderately divergent.

51
The genotypes were grouped in such a way that cluster I had the largest member of all clusters,
included 27 (55%) genotypes, followed by cluster III included 15 (30.6%), cluster IV included 3
(6.04%) genotypes. In contrast cluster V and cluster VI had the smallest member, constituted of 1
(2.04%) genotype each. This cluster analysis revealed that bread wheat genotypes originated from
different sources.

In the present study, genotypes gained from different source center clustered in the same
category together, for instance, in cluster I genotypes released from Adet, Holeta, Mekele,
Kulumsa, and Sinnana grouped together. In support of this Ali et al., 2008; Hailegiorgis et al.,
2011; Fellahi et al., 2013 noted morphological diversity is more important factor rather than
variation in geographical or origin as an indicator of genetic diversity. Moreover, genotypes
collected from the same source of center (Gassay and Tay from Adet) were clustered in to
different clusters, suggesting the existence of genetic diversity within each collection sources.

Table 11. Distribution and grouping of 49 bread wheat genotypes into different
diversity classes of cluster membership based on D 2 analysis

Cluster Number of Name of genotypes Proportions

genotypes (%)
I 27 Mekelle-3, Shorima, ETBW 7368, Hidasse, ETBW 8517, 55.1
ETBW 7120, Sofumar, ETBW 7038, Kakaba, Pavon-76,
ETBW 8512, ETBW 7871, Millan, ETBW 6861, ETBW 8506,
Mekele-4, Biqa, ETBW 8513, ETBW 7058, ETBW 8519,
ETBW 7101, ETBW 8515, Alidoro, Mada-Wolabu, Gassay,
Ogolcho, Dandaa
II 2 Tay, ETBW 7872 4.08
III 15 ETBW 8507, Jeferson, ETBW 8510, Honqolo, Hulluka, Pastor, 30.6
ETBW 7194, ETBW 7364, ETBW 8507, ETBW 8514, ETBW
8509, ETBW 8516, King Bird, ETBW 8518, Hoggana
IV 3 ETBW 8511, ETBW 6940, ETBW 7213 6.12
V 1 Digelu 2.04
VI 1 ETBW 7147 2.04

52
4.11. Cluster Mean Analysis

The mean value of the 11 quantitative characteristic feature are presented in (Table 12). Cluster I
had a characteristics feature of short in days to heading, high values in terms grain yield per
hectare, and moderate high values in terms of harvest index. Cluster II had a characteristics
feature of short in days to maturity, high values in terms of plant height, spike length, number of
spike lets per spike, biological yield and grain yield, while relatively low in harvest index as
compared to other clusters. Cluster III showed short in days to heading and days to maturity as
well as grain yield.

Cluster IV had a characteristics feature of relatively low values of biological yield, and grain
yield per hectare and relatively moderate values in terms of characters studied. Cluster V had a
characteristics feature of long in days to heading and days to maturity, high values in terms of
number of grains per spike, thousand seed weight, and harvest index. On the other hand had low
value in terms of spike length, biological yield, and grain yield per hectare. Cluster VI had a
characteristics feature of long in days to heading and days to maturity, short in plant height,
moderate high values in terms of spike length, and thousand seed weight, and also characterized
by high harvest index and grain yield per hectare.

Therefore, as presented in the table below low and high mean value recorded between cluster (I,
II, III) and cluster (V, VI) for days to heading, between (II, III, V) and (VI) for days to maturity,
between VI and II for plant height, cluster VI and I for number of productive tillers per plant,
cluster V and II for spike length, cluster VI and II for number of spikelets per spike, cluster VI
and V for number of grains per spike, cluster IV and VI for thousand seed weight, cluster V and
II for biological yield, and for harvest index observed between cluster II and VI respectively. In
addition to these the highest grain yield obtained from cluster II, II, VI, and the low grain yield
obtained from cluster III, IV, and V.

53
Table 12. Mean values of seven clusters for 11 characters of 49 bread wheat genotypes

Cluster Means
Cluster HD MD PH NPTP SL NSPS NGS TSW BY HI GY
I 65.50 127.33 81.22 1.55 7.69 14.64 38.14 41.83 2.16 0.326 36.02
II 66.38 126.15 91.8 1.45 8.75 16.3 41.25 41.15 2.48 0.285 36.31
III 65.87 126.41 72.40 1.45 7.07 14.24 36.43 39.29 2.07 0.301 32.51
IV 71.33 132.67 75.73 1.48 8.07 14.93 39.60 38.50 2.00 0.300 30.96
V 79.50 136.30 80.9 1.45 6.80 15.00 45.70 43.10 1.78 0.330 31.75
VI 79.00 136.50 69.2 1.30 7.20 13.30 29.10 43.30 2.05 0.340 36.00
NB: DH=days to heading, DM=days to maturity, PH=plant height, NPTP=number of
productive tillers per plant, SL =spike length, NSPS=number of spikelets per spike,
NGS=Number of grains per spike, BY=biological yield, HI=harvest index, TSW=
thousand seed weight, and GY= grain yield.

54
 5. SUMMARY AND CONCLUSION
Overall variability within a crop is due to heritable and non-heritable components. The present
study comprised 49 bread wheat genotypes that were evaluated at Jamma and Geregera
environments with the overall objective of studying genetic variation and character associations
for 12 traits. The analysis of variance revealed highly significant differences at (P< 0.01 and
p<0.05) levels among the genotypes for all traits except grain filling period at Geregera, which
indicated the existence of variation among the tested genotypes.

Maximum values of genotypic coefficient of variation were recorded for spike length (8.66%),
followed by number of productive tillers per plot (8.4%), number of grains per spike (6.4%) and
thousand seed weight (6.15%), whereas higher values of phenotypic coefficient of variation were
recorded for productive tillers followed by grain yield, spike length and harvest index with a
values of (13.3%, 11.35%, 10.3%, 9%), respectively. Heritability ranged from 29.1% for grain
yield to 82 % for heading date. Relatively high genetic advance as percent of the mean were
recorded for spike length followed by number of productive tillers per plant, number of grains
per spike, thousand-seed weight, days to heading and plant height with values of (14.9%, 10.6%,
10%, 10%, 9.7%, and 9.07%), respectively.

Because of high genotype-environment (G × E) interactions, estimates of GCV, H2 and GA for

most of the characteristics using combined over location analysis were generally lower than the
estimates computed from the variance analyses made separately for each location.

Grain yield displayed positive and significant association with plant height, number of tillers per
plant, thousand seed weight, biological yield, and harvest index at genotypic and phenotypic
level. The estimated ranges of mean values revealed that bread wheat genotypes reflect good
amount of genetic variability, and out of the 49 genotypes, 12 released varieties and 12 pipelines
were characterized by relatively better yield performance as each scored above the overall mean
of 34.6 qt/ha.

Based on the results of the individual and combined analysis of variance, high estimates of
genotypic coefficient of variation, heritability and genetic advance as percent of mean observed
for spike length and thousand seed weight.

55
The principal component analysis revealed that five principal components, with Eigen values
greater than unity, explained 80.4% of the total variability, and hence, grain yield, biological
yield, number of grains per spike, harvest index, and thousand seed weight were the major
contributing characters for variability contained in the bread wheat genotypes.

The highest inter-cluster distance was exhibited between cluster I and III (D2 = 25.79**),
followed by cluster II and IV (D2 = 22.82), cluster II and III (D2 = 22.75), indicating wider
genetic among the clusters. Therefore, initiating crossing program between members of cluster I
with members of cluster III, and members of cluster II with members of cluster III, and IV may
produce a high amount of heterotic expression in the F1’s and broad spectrum of variability in
segregating (F2) populations.

The genetic parameters of the present study revealed that plant height, spike length, number of
grains per spike, and thousand seed weight showed moderate to high heritability and genetic
advance in percentage of mean. High significant positive correlation along with maximum
positive direct effects on grain yield were achieved for harvest index and biological yield, may
be identified as a best selection criterion (trait) for the development of modern wheat variety.

Thus, the results suggest that plant height, higher number of grains per spikes, thousand seed
weight (bold size grains), biological yield and higher harvest index are the important yield
contributing traits and thus plant selection based on these traits will be most effective for future
wheat yield improvement program.

56
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 7. APPENDIX

Appendix Table 1. Mean value of 11 quantitative traits of 49 tested bread wheat

genotypes

SN Genotype DH DM PH NPTP SL NSPP NGS TSW BY HI GY

1 Mekelle-3 63.50 124.0 83.8 1.45 6.7 14.7 40.9 37.3 2.20 0.355 40.00

2 Mada-Wolabu 67.25 127.0 82.1 1.68 8.0 14.1 38.6 44.6 2.35 0.353 43.00

3 ETBW 8508 64.75 125.2 69.5 1.25 7.0 14.6 36.9 36.1 2.13 0.278 30.63

4 Shorima 67.25 128.2 78.3 1.25 7.3 14.5 35.4 39.9 1.93 0.320 31.38

5 Danda'a 70.00 133.3 83.8 1.7 6.9 14.0 41.6 44.9 2.23 0.328 37.50

6 ETBW 7871 67.00 129.5 79.1 1.5 8.2 15.3 40.1 39.3 2.20 0.323 37.50

7 ETBW 8517 64.00 127.3 78.5 1.25 8.2 13.6 35.6 40.4 1.95 0.358 33.75

8 ETBW 6861 70.00 131.0 78.7 1.3 8.4 15.5 35.7 39.2 2.05 0.345 36.25

9 ETBW 7120 66.75 127.3 82.7 1.55 8.3 15.1 36.9 40.2 2.05 0.283 32.50

10 MILLAN 68.00 128.5 79.3 1.25 7.8 13.9 37.2 39.7 2.45 0.313 39.88

11 ETBW 8506 63.00 125.5 77.7 1.45 7.3 14.1 35.6 42.1 2.13 0.335 38.25

12 Mekelle-4 66.00 125.8 79.1 1.25 7.9 14.3 37.8 43.9 2.13 0.345 37.50

13 ETBW 7147 79.00 136.5 69.2 1.3 7.2 13.3 29.1 43.3 2.05 0.335 36.00

14 ETBW 8510 65.25 125.2 71.4 1.25 7.3 14.7 33.9 38.1 2.05 0.290 32.13

15 TAY 67.50 126.5 89.8 1.5 8.0 16.3 40.3 39.2 2.45 0.298 37.63

16 Sofumar 64.00 126.7 83.6 1.7 8.0 16.1 35.6 40.2 2.08 0.310 32.88

17 ETBW 7038 61.25 126.5 78.1 1.65 7.2 14.6 41.1 37.4 2.03 0.328 35.00

18 Hulluka 65.75 127.7 72.6 1.45 6.8 13.7 39.6 35.9 1.95 0.290 30.00

19 ETBW 8511 70.00 134.0 75.7 1.7 8.1 15.0 37.4 36.1 2.20 0.263 29.63

20 ETBW 7368 65.75 128.0 79.9 1.4 8.3 14.6 36.5 39.7 2.05 0.275 29.25

21 Ogolcho 67.00 125.7 86.0 1.35 7.7 15.1 42.1 48.1 2.25 0.328 38.63

22 ETBW 7872 65.25 125.8 93.8 1.4 9.5 16.3 42.2 43.1 2.50 0.268 35.00

66
23 Digelu 79.50 136.3 80.9 1.45 6.8 15.0 45.7 43.1 1.78 0.330 31.75

24 ETBW 7194 71.25 128.0 70.0 1.3 6.4 13.7 34.4 41.8 2.03 0.278 30.63

25 PASTOR 66.00 126.8 76.2 1.35 6.9 13.9 42.0 34.8 2.13 0.278 30.00

26 Kakaba 62.25 124.7 81.2 1.95 7.4 14.1 36.6 39.0 2.03 0.348 35.25

27 ETBW 7364 67.75 127.0 72.8 1.5 7.4 14.1 36.9 42.9 2.05 0.303 31.88

28 ETBW 8512 63.00 125.0 83.0 1.4 7.3 14.1 42.0 38.3 1.85 0.315 31.75

29 Honqolo 66.50 127.0 73.8 1.72 6.8 13.8 32.4 37.2 2.18 0.300 33.75

30 ETBW 8516 61.50 124.5 74.5 1.4 7.1 14.4 40.3 41.4 1.90 0.352 35.00

31 ETBW 6940 72.25 131.8 75.1 1.4 7.8 13.8 37.0 40.8 1.85 0.328 32.00

32 Hidasse 65.00 127.2 79.8 2.0 7.0 13.3 38.1 42.1 2.10 0.313 31.00

33 ETBW 8513 64.00 128.2 83.4 1.9 7.3 14.1 37.0 47.2 2.25 0.335 38.00

34 ETBW 8515 63.50 126.0 79.9 1.65 7.1 14.1 38.5 46.2 2.10 0.290 32.13

35 ETBW 8514 65.25 125.7 68.1 1.72 6.5 13.1 36.6 42.0 2.08 0.270 29.50

36 ETBW 8509 67.25 128.2 72.9 1.2 7.9 16.9 34.0 39.6 2.05 0.258 26.50

37 Alidoro 69.00 129.3 85.1 1.4 10.3 17.7 36.4 45.2 1.78 0.340 30.63

38 Pavon-76 66.25 126.5 79.2 1.35 7.3 13.8 38.4 37.2 2.43 0.275 34.13

39 ETBW 8519 62.50 126.5 83.3 1.85 7.5 15.0 38.8 43.8 2.05 0.345 37.50

40 Jeferson 63.25 124.3 72.8 1.5 7.1 14.6 35.5 35.3 2.03 0.285 30.88

41 Hoggana 71.50 128.8 72.1 1.6 7.6 14.6 32.6 41.8 2.15 0.325 38.25

42 King Bird 62.50 125.8 73.5 1.5 6.7 13.7 37.9 38.6 2.13 0.335 36.63

43 ETBW 8518 62.25 124.2 70.9 1.55 6.8 14.1 38.5 39.5 2.15 0.343 39.38

44 ETBW 7213 71.75 132.2 76.4 1.35 8.3 16.0 44.4 38.6 1.95 0.313 31.25

45 ETBW 7101 64.75 125.5 84.1 1.55 7.9 15.5 36.8 41.5 2.15 0.323 35.95

46 ETBW 7058 66.75 130.0 81.8 1.55 7.3 14.2 35.3 44.7 2.48 0.303 37.75

47 Biqa 64.00 128.0 77.8 1.9 7.3 14.6 39.8 44.0 2.33 0.350 41.50

48 Gassay 66.75 126.8 83.8 1.7 7.6 15.3 41.5 43.4 2.65 0.325 43.75

49 ETBW 8507 67.25 127.8 74.8 1.4 7.7 13.7 34.9 44.4 2.10 0.318 32.50

67
 Appendix Table 2. Estimated values of mean squares and f values of 49 bread wheat genotypes for
12 traits tested at Jamma, using Simple lattice design

Source of Mean square of characters

variance DF HD MD GFP PHT NPTP SL NSPS NGS TSW BY HI GY
Replication 1 9.2 0.09 2.95 73.6 3.2 24.9 16.7 0.79 28.1 0.007 0.009 140.2
Blocks (rep) 12 2.7 3.34 7.00 27.4 0.07 0.81 1.39 7.93 4.58 0.028 0.0005 20.6
Genotype 48 38.5** 15** 16.2* 67** 0.16* 2.1** 1.93* 21.8*32** 0.063* 0.0026** 64.0**
Intra-b error 36 2.77 2.26 8.24 20.6 0.075 0.66 0.89 6.87 6.66 0.031 0.0008 26.75
RCBD 48 2.75 2.53 7.93 22.3 0.073 0.70 1.01 11.1 6.14 0.030 0.074 25.20
CV (%) 2.47 1.23 7.50 5.70 15.7 10.1 6.36 8.00 5.20 7.08 6.89 10.45
R2 (%) 0.93 0.85 0.67 0.76 0.75 0.79 0.69 0.69 0.84 0.674 0.794 0.73
Mean 67.1 129 62.5 83.0 1.72 8.24 15.9 40.0 48.0 2.470 0.388 48.0
E.R RCBD 99.0 104 96.2 102 97.5 101 104 91.4 92.2 97.00 91.00 94.2
LSD at 1% 4.460 4.03 7.7 12.2 0.73 2.18 2.50 9.30 6.90 0.480 0.075 13.9
LSD at 5% 3.34 3.02 5.8 9.10 0.55 1.63 1.90 7.00 5.20 0.360 0.056 10.4
δ2 g 17.87 6.37 3.98 23.2 0.042 0.72 0.52 7.47 12.7 0.0160 0.0009 18.6
δ2 p 19.25 7.50 8.10 33.5 0.08 1.05 0.97 10.9 16.0 0.0315 0.0013 32.0
GCV (%) 6.30 1.96 3.20 5.80 11.9 10.3 4.54 6.83 7.42 5.120 7.6920 9.00
PCV (%) 6.54 2.12 4.55 6.97 16.5 12.4 6.12 8.25 8.33 7.186 9.2450 11.8
H2 (%) 92.8 84.9 49.0 69.25 53.0 68.6 54.0 68.5 79.2 50.79 69.230 58.0
GA 8.40 0.10 2.88 8.27 0.31 1.45 1.10 4.67 6.54 0.172 0.1860 6.77
GAM 12.52 0.08 4.60 9.96 18.0 17.6 6.90 11.7 13.6 7.034 7.5300 14.1
NB: *, ** Indicates significance at 0.05 and 0.01 probability levels, respectively, σ2g = genotypic variance,
2
σ p=phenotypic variance, GCV (%) = genotypic coefficient of variation, PCV (%) = phenotypic
coefficient of variation, H2 (%) =broad sense heritability, GA =genetic advance, GAM % =genetic
advance as percentage of mean, DF=degrees of freedom, DH=days to heading, DM=days to
maturity, GFP= grain filling period, PH=plant height, NPTP=number of productive tillers per plant,
SL =spike length, NSPS=number of spikelets per spike, NGS=Number of grains per spike,
BY=biological yield, HI=harvest index, TSW= thousand seed weight, and GY= grain yield.

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Appendix Table 3. Estimated values of mean squares and f values of 49 bread wheat genotypes for
12 traits tested at Geregera, using Simple lattice design

Source of Mean square of characters

variance DF HD MD GFP PHT NPTP SL NSPS NGS TSW BY HI GY
Replication 1 1.72 1.47 2.90 624 0.276 1.47 1.62 4.76 2.30 0.037 0.002 11.1
Blocks (rep) 12 3.40 9.10 10.5 27.0 0.014 0.15 1.00 6.74 10.3 0.044 0.0008 16.0
Genotype 48 30** 32** 18.6ns 95** 0.11* 1.0** 2.7* 33** 26* 0.165* 0.0027* 41.42*
Intra-b error 36 3.33 9.34 12.5 29.3 0.06 0.23 1.20 13.0 9.0 0.066 0.0012 21.02
RCBD 48 3.47 9.28 12.0 28.8 0.05 0.21 1.24 11.5 9.3 0.060 0.0011 19.78
CV (%) 2.82 2.42 5.77 7.30 17.1 6.76 8.37 7.94 9.0 13.83 14.00 21.00
R2 (%) 0.89 0.77 0.61 0.79 0.70 0.83 0.70 0.80 0.73 0.733 0.708 0.678
Mean 66.0 126 60.0 73.4 1.28 6.82 13.3 35.7 33.8 1.780 0.240 21.17
E.R RCBD 100 99.0 96.0 98.0 81.0 90.7 94.0 88.0 100 91.70 90.40 94.10
LSD at 1% 4.90 8.20 9.7 14.5 0.65 1.30 3.10 9.7 8.00 0.688 0.095 13.30
LSD at 5% 3.67 6.10 7.1 11.0 0.49 0.97 2.30 7.2 6.00 0.516 0.071 9.220
δ2 g 13.34 11.3 0.00 32.9 0.025 0.39 0.75 10.0 8.50 0.050 0.0008 10.21
δ2 p 15.00 16.0 0.00 45.0 0.055 0.50 1.35 16.5 13.0 0.082 0.0013 20.71
GCV (%) 5.530 2.67 0.00 7.80 12.4 9.10 6.51 8.86 8.63 12.56 11.78 15.10
PCV (%) 5.870 3.17 0.00 9.14 18.3 10.4 8.74 11.4 10.7 16.13 15.00 21.50
H2 (%) 88.90 70.8 0.00 73.0 45.5 77.0 55.6 60.6 65.4 60.00 55.55 49.30
GA 7.100 5.84 0.00 10.1 4.02 1.12 1.33 5.08 4.86 0.350 0.041 4.630
GAM 10.76 4.50 0.00 13.8 31.4 16.5 10.0 14.2 14.4 20.00 17.21 21.86
NB: *, ** Indicates significance at 0.05 and 0.01 probability levels, respectively, σ2g = genotypic variance,
2
σ p=phenotypic variance, GCV (%) = genotypic coefficient of variation, PCV (%) = phenotypic
coefficient of variation, H2 (%) =broad sense heritability, GA =genetic advance, GAM % =genetic
advance as percentage of mean, DF=degrees of freedom, DH=days to heading, DM=days to
maturity, GFP= grain filling period, PH=plant height, NPTP=number of productive tillers per plant,
SL =spike length, NSPS=number of spikelets per spike, NGS=Number of grains per spike,
BY=biological yield, HI=harvest index, TSW= thousand seed weight, and GY= grain yield.

69
 Alidoro

4.0

3.2 Digelu
ETBW 7213
2.4
Component 2 (20.58%)

SL ETBW 7872
ETBW 8509 Hd Md NSPP
ETBW 6861
1.6 ETBW 7147 ETBW 8511
ETBW 6940
ETBW 7120 TAY
PHT
0.8 ETBW 7368 ETBW 7871
TSW
Sofumar Danda'a Ogolcho
Shorima
0.0 Hoggana NGS
ETBW 7101
ETBW 8507
ETBW 8517 Mekelle-4
MILLAN
ETBW 7194 ETBW 7364 ETBW 7058
ETBW 8510 HI Mada-WolabuGassay
-0.8 PASTOR ETBW 8512
Pavon-76 BYP ETBW 8519
ETBW 8508
Hulluka ETBW 8515 GYP 8513
ETBW
Jeferson ETBWETBW
85167038 8506 NPT
ETBW Biqa
-1.6 Hidasse
Honqolo Mekelle-3
Kakaba
ETBW 8514 King Bird
-2.4
ETBW 8518

-2.4 -1.6 -0.8 0.0 0.8 1.6 2.4 3.2 4.0

Component 1 (25.48%)

Appendix Figure 1. Principal components plot of bread wheat genotypes based on 11

agronomic and phenotypic traits.

70
Appendix Figure 2. Tree diagram of genetic relationships among 49 bread wheat
genotypes.

71