BAHIR DAR UNIVERSITY

COLLEGE OF AGRICUTLURE AND ENVIRONMENTAL SCIENCES

GRADUATE PROGRAM

GENETIC DIVERSITY AND ASSOCIATION OF TRAITS IN ETHIOPIAN FABA BEAN

(Vicia faba L.) ACCESSIONS

M.Sc. Thesis

By

Andualem Muche

August 2020
Bahir Dar, Ethiopia
 BAHIR DAR UNIVERSITY
COLLEGE OF AGRICUTLURE AND ENVIRONMENTAL SCIENCES
GRADUATE PROGRAM

GENETIC DIVERSITY AND ASSOCIATION OF TRAITS IN ETHIOPIAN FABA BEAN

(Vicia faba L.) ACCESSIONS

M.Sc. Thesis

By

Andualem Muche

SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE

DEGREE OF MASTER OF SCIENCE (M.Sc.) IN PLANT BREEDING

August 2020
Bahir Dar, Ethiopia
 THESIS APPROVAL SHEET

As member of the Board of Examiners of the Master of Science (M.Sc.) thesis open defense
examination, we have read and evaluated this thesis prepared by Mr. Andualem Muche,
entitled with “Genetic diversity and association of traits in Ethiopian faba bean (Vicia
faba L.) accessions”. We hereby certify that; the thesis is accepted for fulfilling the
requirements for the award of the degree Master of Science (M.Sc.) in Plant Breeding.

Board of Examiners

Mulugeta Atnaf (PhD) -------------

Name of External Examiner Signature Date

Tiegist Dejene (PhD) ------------------

Name of Internal Examiner Signature Date

Wossen Tarekegne (PhD) ---------------- ------------------

Name of Chair Person Signature Date
 DECLARATION

This is to certify that, this thesis entitled with “Genetic diversity and association of traits in
Ethiopian faba bean (Vicia faba L.) accessions” submitted in partial fulfillment of the
requirements for the award of the degree of Master of Science (M.Sc.) in Plant breeding to
the Graduate Program of College of Agriculture and Environmental Sciences, Bahir Dar
University by Mr. Andualem Muche (ID. No. BDU1100586PR) is an authentic work carried
out by him under our guidance. The matter embodied in this work has not been submitted
earlier for award of any degree or diploma to the best of our knowledge and belief.

Name of the Student

Andualem Muche
Signature ____________date ______________

Name of the Advisors

1) Alemu Abate (PhD) (Major Advisor)
Signature ____________date ______________

2) Fisseha Worede (PhD) (Co- Advisor)

Signature ____________ date ______________
 ACKNOWLEDGEMENT

First of all, I am highly grateful to The Almighty God who gave me good health, guidance,
protection and abundant blessings throughout my studies and because of You; I made this
through against all odds.

I would like to express my special gratitude and heartfelt thanks to my major supervisor, Dr.
Alemu Abate, for his practical and technical guidance, constructive comments, suggestions
and encouraging the use of correct grammar and consistent notation in my writings and for
carefully reading and commenting on countless revisions of this manuscript assistance from
the very beginning to the completion of this thesis write-up. I would also like to extend my
special gratitude to my co-supervisor, Dr. Fisseha Worede, for his keen interest in supporting
my research comments, suggestion and for contributing towards improving the manuscript.

I would like to thank Bahir Dar University, college of Agriculture and environmental sciences
for providing me opportunity to study my M.Sc. degree and experimental plot for the field
research. I would also like to thank Ethiopian biodiversity institute for the provision of
accessions for the study.

I would like also to thank all my classmates for their essential support and giving me
necessary information, especially, Mr. Abunu Marefia and Asaye Birhanu assisted me with
inestimable technical and moral support in addition to providing me with the help to run
different soft ware to conduct the data analysis starting from the very beginning in the field up
to the completion of this thesis write-up.

Particular gratitude and appreciation goes to my father Muche Hiywotu and my beloved
mother Gedam Addis whose crucial decision to invest part of their meager resource in my
education and all my family members for bringing me up with their priceless love, care, for
encouraging and supporting me. I also express my heartfelt thanks to my brothers; Tilahun
Muche, Yalew Muche, Habtamu Muche and all my sisters for their financial and moral
support specially my brother Tilahun Muche dreams and determination to educate me until I
reached M.Sc. level have come true and he had laid a base for me to reach where I am today. I
also express my heartfelt thanks to all my friends for their marvelous support during this
research work.
 ABBREVIATIONS AND ACCRONYMS

ANOVA Analysis of Variance

CSA Central Statistical Authority
CV Coefficient of Variation
FAO Food and Agriculture Organization
FAOSTAT Food and Agriculture Organization Statistics
GA Genetic Advance
GAM Genetic Advance as percent of Mean
GCV Genotypic Coefficient of Variation
ICARDA International Center for Agricultural Research in Dry Areas
MT Metric ton

PC Principal Component

PCA Principal Component Analysis

PCV Phenotypic Coefficient of Variation
SAS Statistical Analysis System
SD Standard Deviation
UPGMA Unweighted Pair Group Method with Arithmetic Averages
 GENETIC DIVERSITY AND ASSOCIATION OF TRAITS IN ETHIOPIAN FABA
BEAN (Vicia faba L.) ACCESSIONS

By: Andualem Muche

Advisors: Alemu Abate (PhD) and Fisseha Worede (PhD)

ABSTRACT

Knowledge on genetic diversity and association of traits in faba bean (Vicia faba L.) is
crucial to enhance productivity and resistance for biotic and abiotic factors. However, there
has been very limited information on variability and association of traits in faba bean
accessions. An investigation was conducted to study phenotypic and genotypic variability,
heritability and genetic advance of yield and associated traits of faba bean accessions at
Kudmi in 2019 main cropping season. The total experimental materials were 81 faba bean
accessions. The design was 9 X 9 simple lattice. The analysis of variance result showed that
highly significant (p<0.01) differences among accessions were observed for all traits except
days to maturity and number of seeds per pod. High phenotypic and genotypic coefficient of
variation was observed for 100 seed weight, biomass yield, harvest index, grain yield, number
of branches per plant, number of pods per plant, ascochyta blight, chocolate spot and rust.
High broad sense heritability estimates were found for traits such as days to flowering
number of pods per plant, biomass yield, grain yield, 100 seed weight, harvest index,
ascochyta blight, chocolate spot and rust disease score. High GAM was recorded for number
of pods per plant, biomass yield, 100 seed weight, number of branches per plant, grain yield,
harvest index, ascochyta blight, chocolate spot and rust disease score. The result of
correlation analysis revealed that grain yield had highly significant (p<0.01) and positive
phenotypic and genotypic correlations with plant height, pod length, number of pods per
plant, number of branches per plant, biomass yield, 100 seed weight and harvest index. High
positive direct effects on grain yield were exerted by biological yield and harvest index at
phenotypic and genotypic levels. Cluster analysis distinguished the 81 accessions into seven
clusters. The first four principal components (eigen values > 1) explained 73.81% of the total
variations among the 81 faba bean accessions. Generally, it can be concluded that there was
considerable variations among faba bean accessions for the traits considered, implying the
possibility of improving these traits through direct and indirect selection. These genetic
resources should be further characterized over seasons and locations and using molecular
tools.

Keywords: Clustering, Coefficient of variance, Correlation, Faba bean, Genetic advance,

Heritability, Path analysis, Variability
 TABLE OF CONTENTS

Contents Pages
THESIS APPROVAL SHEET ii
DECLARATION iii
ACKNOWLEDGEMENT iv
ABBREVIATIONS AND ACCRONYMS v
ABSTRACT vi
LIST OF TABLES x
LIST OF FIGURES xi
LIST OF APPENDICES xii
Chapter 1. INTRODUCTION 1
1.1. Background and Justification 1
1.2. Statement of the Problem 2
1.3. Objective of the Study 3
1.3.1. General objective 3
1.3.2. Specific objectives 3
Chapter 2. LITERATURE REVIEW 4
2.1. Origin and Domestication of Faba Bean 4
2.2. Botanical Characterization 4
2.3. Agro ecology and Distribution in Ethiopia 5
2.4. Faba Bean Production in Ethiopia 6
2.5. Faba Bean Production Constraints in Ethiopia 7
2.6. Importance of Faba Bean 8
2.6.1. Consumption 8
2.6.2. Nutritional content as food 8
2.6.3. Source of nutrient for soils 9
2.7. Faba Bean Genetic Diversity 9
2.8. Genetic Variability 11
2.9. Heritability 12
2.10. Genetic Advance 14
 TABLE OF CONTENTS (Continued)

2.11. Correlation of Traits 15

2.12. Path Coefficient Analysis 17
2.13. Cluster Analysis 18
2.14. Principal Component Analysis 19
Chapter 3. MATERIALS AND METHODS 21
3.1. Description of the Study Area 21
3.2. Experimental Materials and Design 21
3.3. Data Collected 21
3.3.1. Data collected on plant basis 22
3.3.2. Data collected on plot basis 24
3.4. Data Analyses 24
3.4.1. Estimation of variance components 26
3.4.2. Correlation analysis 28
3.4.3. Path coefficient analysis 29
3.4.4. Clustering of accessions and genetic divergence analysis 30
3.4.5. Principal component analysis 31
Chapter 4. RESULTS AND DISCUSSION 32
4.1. Variability among Accessions 32
4.2. Estimation of Mean and Range 33
4.3. Estimates of Genetic Parameters 38
4.3.1. Phenotypic and genotypic coefficients of variation 38
4.3.2. Broad sense heritability 40
4.3.3. Genetic advance 41
4.4. Correlation among Traits 42
4.4.1. Phenotypic correlation of grain yield with other traits 42
4.4.2. Phenotypic correlation among other traits 43
4.4.3. Genotypic correlation of grain yield with other traits 45
4.4.4. Genotypic correlation among other traits 45
4.5. Direct and Indirect Effects of Traits 48
 TABLE OF CONTENTS (Continued)

4.5.1. Phenotypic path coefficients 48

4.5.2. Genotypic path coefficients 49
4.6. Clustering among Accessions 53
4.6.1. Genetic distance among clusters 56
4.6.2. Mean performance of clusters 57
4.7. Principal Component Analysis 59
Chapter 5. CONCLUSION AND RECOMMENDATIONS 62
5.1. Conclusion 62
5.2. Recommendations 63
6. REFERENCES 64
7. APPENDICES 84
BIOGRAPHICAL SKETCH 95
 LIST OF TABLES

Tabl Page
e
3.1. Rating scale of diseases 23
3.2. The structure of analysis of variance (ANOVA) for simple lattice design 25
4.1. Analysis of variance for 14 traits of eighty-one faba bean accessions 32
4.2. Minimum, mean and maximum values of 14 traits of 81 faba bean accessions 35
4.3. Extent of variability in qualitative traits among faba bean accessions 37
4.4. Estimates of variances, coefficient of variation, heritability and genetic
advance of traits 39
4.5. Estimates of genotypic (below diagonal) and phenotypic (above diagonal)
correlation coefficients of 12 traits of 81 faba bean accessions 47
4.6. Phenotypic direct (bold and diagonal) and indirect effects of 11 traits on grain
yield of 81 faba bean accessions 51
4.7. Genotypic direct (bold and diagonal) and indirect effects of 11 traits on grain
yield for 81 faba bean accessions 52
4.8. List of faba bean accessions grouped into seven clusters 54
4.9. Estimates of average intra (diagonal) and inter (above diagonal) cluster
distances for 7 clusters of 81 faba bean accessions 57
4.10. Cluster means for 12 traits of 81 faba bean accessions 58
4.11. Eigenvectors, eigen values and variance explained by the first four principal
components 60

LIST OF FIGURES
Figure Page
4.1. Diagrammatic representation of qualitative traits observed in 81 Faba bean
accessions 37
4.2. Dendrogram showing relationships among 81 Ethiopian faba bean
accessions using 55
4.3. Principal component analysis biplot describing the relative position of 81
faba bean accessions and 12 traits 61
 LIST OF APPENDICES

Appendix Page
Table 1. List of the 81 faba bean accessions used in the study 85
Table 2. Mean performance of 81 Faba bean accessions for 14 traits tested at
Kudmi in 2019 89
 Chapter 1. INTRODUCTION

1.1. Background and Justification

Faba bean (Vicia faba L.), 2n = 2x = 12 is one of the earliest domesticated food legumes in
the world (Singh et al., 2013). Faba bean belongs to the family Fabaceae, sub-family
Faboideae, tribe Fabeae, and the genus Vicia. Vicia has a large genus of 140-190 species
(Doyle and Luckow, 2003). Faba bean is one of the most important legume crops and is
believed to have originated in the Near East and cultivation started early in Neolithic times,
8000 B.C. (Cubero, 1974; Torres et al., 2006; Karkanis et al., 2018).

Due to its high nutritional value and its ability to grow over a wide range of climatic and soil
conditions (Yahia et al., 2012), faba bean is the fourth most important legume worldwide,
after pea, chickpea and lentil. It is grown under different cropping systems for its dry grains,
green pods, animal feed and a green-manure in the world. The major faba bean growing
countries are China (1.80 Mt), Ethiopia (0.93 Mt), Australia (0.37 Mt), Germany (0.19 Mt),
France (0.19 Mt), Egypt (0.11 Mt) and Sudan (0.11 Mt) (FAOSTAT, 2019). China is the
leading producer with 37% share of the world’s total faba bean production (FAOSTAT,
2019).

Faba bean is one of the major pulse crops grown in the highlands (1800-3000 m.a.s.l) of
Ethiopia (Temesgen Alene and Aemiro Bezabih, 2012; Tafere Mulualem et al., 2012).
Ethiopia is the second largest producers of faba bean in the world, next to China. However,
the national productivity of faba bean in the country is still very low. According to the report
of Central Statistics Agency (CSA), the national average yield of faba bean is about 2.1 t ha-1
(CSA, 2018) which is very low compared to the average yield of 3.7 t ha -1 in major producer
countries (FAOSTAT, 2017).

Faba beans need a cool season for best growth with moderate amounts of rainfall 650 to 1000
mm per annum (Gasim and Link, 2007) and considered to tolerate frost, but are susceptible to
drought and water logging (Subash and Priya, 2012). Faba beans are moderately tolerant to
acid soil conditions than most legumes (Singh et al., 2010), but grow best in loamy soils
(Rajan et al., 2012). The crop takes about four to five months for the pods to mature enough
for the seeds to be harvested based on environmental condition (Mussa Jarsso et al., 2008). In
Ethiopia, the major faba bean producing regions are Oromia (Wollega, Shoa and Bale),
Amhara (Gondar, Gojam and Wollo), Southern parts of the country (Gamo Gofa),
Benishangul-Gumuz and highland parts of Tigray (CSA, 2017).

The crop has a potential to substitute meat in many parts of the world where there is demand
for non-animal protein sources (Crepon et al., 2010). Ethiopian farmers are also aware of the
role of the crop in improving soil health by fixing atmospheric nitrogen and widely use it in
rotation with cereals. Faba bean is a source of cash to the farmers and foreign currency to the
country. Despite its diverse benefits, the productivity of the crop is attributed by biotic and
abiotic factors. However, research on this crop was started in 1950’s to tackle different
production constraints of the crop with the prime objectives to improve grain yield, seed size
and resistance to important diseases, particularly chocolate spot (Botrytis fabae) (Gemechu
Keneni et al., 2006).

1.2. Statement of the Problem

Despite its importance, the productivity of faba bean in Ethiopia is constrained by different
factors which include biotic (diseases, weeds and insects), abiotic (drought, water logging and
marginal management) and edaphic stresses like soil acidity. The measures to be done to
confront this challenge are developing varieties resistant to major biotic, abiotic and edaphic
constraints. However, information on variability and association among traits is limited for
Ethiopian faba bean accessions. Moreover, limited attention has been given to the crop to
study phenotypic and genotypic variability, heritability and genetic advance of yield and
associated traits of faba bean accessions in the study areas. In addition, shortage of improved
varieties and limited efforts to improve the crop are major bottle necks for extensive
production of faba bean in the study area. Knowledge on the extent and pattern of genetic
variability present in a population is absolutely essential for further improvement of the crop.
Similarly, information on the extent and nature of interrelationship among trait help in
formulating efficient scheme of multiple trait selection. Besides, knowledge of the naturally
occurring diversity in a population helps to identify diverse groups of accessions that can be
useful for future breeding program. Such information is limited in faba bean production
system of Ethiopia. Therefore, the present study was initiated with the following objectives:

1.3. Objective of the Study

1.3.1. General objective

The overall objective was to assess the extent of genetic diversity and association among
grain yield and yield-related traits of Ethiopian faba bean accessions.

1.3.2. Specific objectives

To evaluate the extent and pattern of genetic diversity among faba bean accessions
To estimate variability, heritability and genetic advance of traits
To estimate association among yield and yield related traits
To group accessions based on their similarity and dissimilarity
 Chapter 2. LITERATURE REVIEW

2.1. Origin and Domestication of Faba Bean

The genus Vicia L. belongs to the family Fabaceae. Knowledge of the wild progenitor and
area of origin of the genus, and subsequent steps in the domestication of its most important
member species, V. faba L., is scarce and disputed (Shiran et al., 2014). The Near East is
considered a center of origin for faba bean (Cubero, 1974), while China seems to be a
secondary center of faba bean genetic diversity (Zong et al., 2009, 2010). In support of
Cubero’s findings, Caracuta et al. (2016) have identified seeds of a potential ancestor of faba
bean adjacent to Mount Carmel, Israel – the remains were C-dated to 14,000 years BP (before
present). Moreover, Caracuta et al. (2015) have determined that faba bean was already
domesticated about 10,200 years BP in the Lower Galilee, Israel. In any case, faba bean can
be considered one of the earliest domesticated crops in light of numerous archeological
findings in Eurasia and Africa which date back to the early Neolithic (Duc et al., 2015).

Vicia faba has a large genetic diversity. According to Duc et al. (2010, 2015), >38,000
accessions of faba bean germplasm are conserved globally in numerous gene banks, as well as
at the International Center for Agricultural Research in Dry Areas (ICARDA). Faba bean has
a diploid (2n) chromosome number of 12, meaning that each cell in the plant has 12
chromosomes (6 homologous pairs). Five pairs are acrocentric chromosomes and 1 pair is
metacentric (Alghamdi, 2009). The wild progenitor and the exact origin of Faba bean remain
unknown. Several wild species (Vicianar bonensis L. and V. galilaea) are taxonomically
closely related to the cultivated crop, but they contain 2n = 14 chromosomes, whereas
cultivated faba bean has 2n = 12 chromosomes. Numerous attempts to cross the wild species
to cultivate faba bean have failed (Hajjar and Hodgkin, 2007).

2.2. Botanical Characterization

Faba bean is a cool season annual legume (Bilalis et al., 2003) that forms coarse, upright,
hollow, and grows between 0.1 and 2 m tall (Duc et al., 2015; Heuze et al., 2016). Stem
growth is indeterminate, and some cultivars are prone to lodging. The leaves are alternate,
pinnate, and consist of two to six leaflets, which are up to 8 cm long without tendrils (Bond et
al., 1985). The flowers have a typically papilionaceous structure and are grouped in
inflorescences; they are either pure white in color or with diffuse anthocyanin pigmentation
on all petals, while black spots are often present on the wing petals (Duc et al., 2015; Heuze et
al., 2016). Seeds, which vary considerably in size, are oblong to broadly oval with a
prominent hilum at the end; their color can be yellow, green, brown, black, or violet, and
sometimes seeds are spotted (Duc et al., 2015). Faba bean plants feature a robust taproot with
frequent lateral root branching from the top of the tap root; nitrogen-fixing nodules containing
rhizobia occur on both the tap and lateral roots (Bond et al., 1985).

Faba bean is generally considered day-neutral, while some accessions require long-day
conditions in order to flower. However, thermal time is the most important contributor to
flowering progress in faba bean, with approximately 830–1000 days above 0°C being
required; winter faba bean genotypes require vernalization (Patrick and Stoddard, 2010). In a
recent study, Cao et al. (2017) found that several potential regulators are implicated in the
vernalization process in faba bean. Faba bean is a self-pollinated plant with significant levels
of cross-pollination (Chen, 2009). The main pollinating insects are honeybees (Apis spp.) and
bumblebees (Bombus sp.); the benefits of insect pollination for yield have been well
documented (Cunningham and Le Feuvre, 2013; Bishop et al., 2016). Faba bean has a diploid
(2n) chromosome number of 12, meaning that each cell in the plant has 12 chromosomes (6
homologous pairs). Five pairs are acrocentric chromosomes and 1 pair is metacentric
(Alghamdi, 2009).

2.3. Agro ecology and Distribution in Ethiopia

Faba bean is one of the major pulse crops grown in the highlands (1800–3000 m.a.s.l) of
Ethiopia (Temesgen Alene and Aemiro Bezabih, 2012; Tafere Mulualem et al., 2012). Faba
bean is an annual herb legume grows best under cool, moist conditions with hollow upright
stems up to 2 m tall. Hot, dry weather is injurious to the crops, especially during flowering
stage, so early planting is important. Faba bean tolerates frost, and rainfall of 650 to 1000 mm
per annum evenly distributed is ideal for the crop (Gasim and Link, 2007). Medium textured
soils with pH ranging from neutral to alkaline (pH of 6.5 to 8.0) (Rajan et al., 2012), are
ideally suited for faba bean production, but it grows best on rich clay loams.

Faba bean is produced in many regions of Ethiopia. The major producing regions are Oromia
(Wollega, Shoa, Bale and Arsi), Amhara (Gondar, Gojam and Wollo), South People Nation
and Nationality, some parts of Tigray and Benishangul-Gumuz. In addition, it is grown in the
pockets in the rest of the country’s high land and semi-highland regions with altitudes ranging
from 1800 to 3000 m above sea level (Tewodros Tesfaye et al., 2015). The total area under
cultivation is estimated to be 3.40% (about 427,696.80 hectares) and the total production is
3.02% (about 8,780,108.79 kilograms (CSA, 2017).

2.4. Faba Bean Production in Ethiopia

Ethiopia is the second largest producer of faba bean in the world after China (Mussa Jarso and
Gemechu Keneni, 2006; FAOSTAT, 2017) through its contribution was not more than 14.7 %
of the amount that the top ten growing countries can contribute (FAOSTAT, 2017).
According to CSA (2016), faba beans are important pulses covering 3.56% (about 443,966.09
hectares) of the grain crop area. The report also revealed that the production obtained from
faba beans, was 3.18% (about 848654.57 ton) of the grain production. While faba bean was
grownup throughout the country, production is targeted within the Amhara and Oromia
regions (Shahidur et al., 2010; Esther, 2013; John et al., 2014; Boere et al., 2015). Amhara
region alone accounts for more than 34% of area coverage and about 31% total volume of
production of the country (CSA, 2018). The primary producers are smallholders with small
and dispersed plots under rain-fed conditions. More than 3,568,225 farmers were involved in
growing faba bean. North Shewa (Amhara region), North Shewa and Arsi zones of Oromia
region are identified as major production areas of faba bean (John et al., 2014; CSA, 2016).

Recently, more than 1.6 million hectares of land has been cultivated to pulses in Ethiopia and
faba bean shares 0.4 million hectares. During the last ten years (between 2007 and 2017), the
area under faba bean cultivation was decreased from 520,519 to 437,106 ha (16.02%) whereas
the production and productivity during the corresponding period was increased from 0.69 to
0.92 million ton (25%) and 1.323 to 2.109 t ha -1(37.27%), respectively (CSA, 2007; 2017).
Faba bean has a versatile use and it is an important source of dietary protein to the majority of
population in Ethiopia (Mussa Jarso and Fasil Assefa, 2014), while its dry seeds, green haulm
and dry straw are used as animal. Trends of productivity and total annual national production
of faba bean between 2005 and 2015 in Ethiopia was progressively increasing despite the
stagnation in cultivated areas as a result of improved productivity per unit area of land
(Gemechu Keneni et al., 2016). The increase in production and productivity of the crop was
due the presence of best technologies regardless of reduction in area of production. Replacing
local varieties with improved ones led to increment in production of 18% in Egypt, 8% in
Sudan and 42% in Ethiopia (ICARDA, 2008). Studies on genetic progresses from breeding of
faba bean also confirmed existence of reasonable levels of yield gain over the last three
decades (Tamene Temesgen et al., 2015).

The productivity of faba bean is in an increasing trend due to the use of improved
technologies. In relation to the desire, however, it is not possible to say that the technologies
developed and promoted through the efforts yet have been readily accepted, fully utilized and
modernized production at farm levels (Gemechu Keneni and Imtiaz, 2010). Though Ethiopian
government is planned to increase faba bean productivity by 32.50% in 2030 (Gemechu
Keneni et al., 2016). However, there is still a long way to go in terms of technical capacity
and capability in order to exploit the high potential of this crop. Periodic assessment of the
previous research achievements and understanding the amount of progresses realized through
research for development efforts enables to improve the efficiency and effectiveness of future
research endeavors (Gemechu Keneni et al., 2016).

2.5. Faba Bean Production Constraints in Ethiopia

In Ethiopia, the productivity of pulses in general and faba bean in particular is still, far below
its potential. Major production constraints includes lack of improved seeds and a high use of
local varieties (Shahidur et al., 2010; Boere et al., 2015), small farm management practices
and land fragmentation. Low soil fertility in the high potential areas is another problem, while
fertilizer use on legume crops is usually low (Solomon Asfaw and Bekele Shiferaw, 2009).
Besides, the crop is threatening by new gall forming disease (Endale Hailu et al., 2014). Very
recently, it was found that the most devastating and widely disseminated in the high land
potential areas within a few years since its occurrence (Endale Hailu et al., 2014).

2.6. Importance of Faba Bean

2.6.1. Consumption

Cultivated faba bean is used as human food in developing countries and as animal feed,
mainly for pigs, horses, poultry and pigeons in industrialized countries (Singh et al., 2012). It
can be used as a vegetable, green or dried, fresh or canned (Gasim and Link, 2007), and as
common breakfast food in Ethiopia, China, Middle East, and Mediterranean countries (Singh
et al., 2013). Faba bean is tightly coupled with every Ethiopian life. It is mainly used as an
alternative with peas to prepare flour called shiro which is used to make shiro wot (a cook
almost ever-present in Ethiopian dishes). The growing importance of faba bean as an export
crop in Ethiopia has crystal rectifier to a revived interest by farmers to extend the area under
production (Samuel Sahile et al., 2008). Ethiopia has been generating around US$30 million
per year from faba bean exports (John et al., 2014).

2.6.2. Nutritional content as food

Faba bean is an excellent source of protein and starch with protein content ranging between
22.0 to 38.2% of seed dry matter, depending on genotype. Therefore, it is considered as a
cheap protein in many developing countries where people cannot afford to buy meat
(Alghamdi et al., 2012). Total carbohydrate content of faba bean is 57.3%, with average
starch content of 47%; faba bean seeds contain 12% fiber and 1.2–4.0% lipids (Baginsky et
al., 2013). Faba bean is also an important source for vitamins and a good source of energy
with 320 kcal/100 g (Ofuya and Akhidue, 2005). Faba bean is a rich source of folates (B
vitamins) with 148 mg/100 g, which have roles in DNA synthesis and repair, amino acid
metabolism and prevention of anaemia by helping in red blood cell production (Singh, 2018).
Faba bean is rich in mineral elements. Karakoy et al. (2018) reported the average nitrogen,
phosphorous, potassium, magnesium, and calcium contents as 6.40, 0.56, 1.51, 0.35, and
0.62%, respectively. According to the same source, faba bean also contains 17.6, 42.7, 83.8,
and 24.0 mg per kg of copper, zinc, iron and manganese, respectively. Faba bean has phenolic
compounds including phenolic acids such as gallic acid, flavonoids, tannins, stillbenes and
lignins. Flavonoids are very important compounds because they have antioxidant, anti-
inflammatory, antiviral, anticancer and anti-atherosclerotic effects (Nijveldt et al., 2001).

2.6.3. Source of nutrient for soils

Several studies have shown that incorporating legume residues into the soil as green manure
increases growth and yield in many crops, such as canola, maize, potato, and wheat (Sincik et
al., 2008; Miller et al., 2011; Bilalis et al., 2012). The amount of nitrogen that can be fixed by
faba bean is mainly dependent on the cultivar, local farming practices (such as nitrogen and
phosphorus fertilization), soil properties, and the presence of symbiotically effective rhizobia
in the soil (Adak and Kibritci, 2016; Anteneh Argaw and Abere Mnalku, 2017). According to
Neugschwandtner et al., (2015), faba bean can fix up to 200 kg N ha -1, while the incorporation
of legume residues into the soil improves soil properties such as organic matter content, bulk
density, porosity, and field capacity (Mandal et al., 2003; Salahin et al., 2013; Adekiya et al.,
2017). Despite its diverse benefits and the availability of high yielding varieties (>3 tha -1)
(MoALR, 2017), in Ethiopia the national average yield of faba bean is about 2.1 t ha -1 (CSA,
2018) has remained low compared to Egypt and United Kingdom 3.47 and 3.83 t ha -1,
respectively (FAOSTAT, 2018).

2.7. Faba Bean Genetic Diversity

The expression “divergence in character” was used by Darwin (1859) for the variation in
genera and species. The knowledge about the distribution and extent of genetic variation
within and between populations (accessions) is essential in order to: estimate any possible loss
of genetic diversity, trace the available genetic variability and its potential in breeding
programmes, select possible population (s) (accession (s)) for in situ and or ex situ
conservation, and show evidence of the evolutionary forces influencing natural populations
(accessions) (Semagn Kassa, 2002). Measurements of genetic diversity were initially based on
co-ancestry and pedigree records (Kim and Ward, 1997).

Primary diversity for the genus is centered in the Near East and Middle East, with a large
percentage of the species occurring in the Irano-Tauranian floristic region and has a wide
morphological diversity. Bond (1976) considered Ethiopia as secondary centers of diversity
for faba bean due to the existence of wide genetic variability. Gemechu Keneni et al. (2003)
indicated the existence of high genetic diversity which was not uniformly distributed among
access and the regions in their study of the extent and pattern of genetic diversity among the
160 faba bean landraces collected from north and south Wello, North Gonder, North Shoa and
Arsi zones. Report by Dawit Tadesse et al. (1994) and Gemechu Keneni et al. (2005) also
revealed the existence of a wealth of genetic variability among Ethiopian faba bean
germplasms for most of the morpho-agronomic traits. Different reports also indicated that
modern plant breeding reduced crop genetic diversity (Vellve, 1993; Tripp, 1996) and thereby
causing to the vulnerability of the crop to biotic and abiotic stresses and failure of their ability
to evolve and adapt to ever changing climates (Gemechu Keneni et al., 2012).

In a species, the pattern of variation can be measured in terms of levels of diversity in

different populations and genetic distances between different populations. Genetic variation
can be attributed to human activities, environmental factors and the dynamic process of
hybridization, segregation and selection. Hence, the variation may be the result of the
interaction between abiotic and biotic factors and population characteristics. Physical and
biotic environmental factors exert selection pressure on variants, leading to the establishment
of specific types, which indicates that the genetic variation is different from place to place
depending on differences in the environment. As regards the biological characteristics of a
species, population structure and mating system determine the level and pattern of genetic
variation. In general, out breeding increases gene flow, resulting in higher polymorphism and
genetic diversity, while selfing restricts gene flow (Hamrick and Godt, 1989). Plant
population heterotic selection maintains out crossing.
Genetic divergence is the statistical distance between the genotypes. It is determined by using
cluster analysis, which assigns genotypes into different groups (Singh and Chaudhary, 1999).
Selection, direct or indirect, can be effective on heritable variability that already exists in a
population. Variability can also be created artificially through hybridization technique by
crossing genetically distant parents. Here comes the importance of genetic divergence
analysis in classifying genotypes into distinct genotypic classes and identifying parents for
hybridization.

Crossing of genotypes belonging to the same cluster would not be expected to yield desirable
recombinants. Consequently, a crossing program might be formulated in such a way that
parents belong to different clusters. The more diverse the parents, within overall limits of
fitness, the greater are the chances of obtaining higher among of heterotic expression of F1’s
and broad spectrum of variability in segregating populations (Rao et al., 1981). The use of D2
statistic (Mahalanobis, 1936) is one of the most important biometrical techniques for
estimating genetic divergences present in a population. Information on the extent of genetic
diversity amongst the breeding materials is very important in the crosses between groups with
maximum genetic divergence would be more responsive for improvement since they are
likely to produce desirable recombination and segregation in their progenies after
hybridization.

2.8. Genetic Variability

Variability is the occurrence of differences among individuals due to differences in their

genetic composition and/or the environment in which they are raised (Allard, 1960; Falconer
and Mackay, 1996). If the character expression of two individuals could be measured in the
environment identical for both, differences in the expression would result from genetic control
and hence such variation is called genetic variation (Welsh, 1981; Falconer and Mackay,
1996). Information on the nature and magnitude of genetic variability present in a crop
species is thus important for developing effective crop improvement program (Singh et al.,
1980; Welsh, 1981).
Phenotypic variability is the total variability which is observable. It includes both genotypic
and environmental variation and hence changes under different environmental conditions.
Such variation is measured in terms of phenotypic variance. In attempting to develop
improved varieties, the plant breeder bases his/her observation often on the measurement of
the phenotype. For plant breeding to be effective there must be phenotypic variation of the
desired trait and some of the variation must be heritable from generation to generation
(Stoskopf et al., 1999). In faba bean, high estimates of variability have been reported for a
number of attributes by Sharifi (2015), Bakhiet et al. (2015), Shelmary and Shivani (2015), Al
Bari and Shtaya (2013), and El-Badawy et al. (2012).

Genotypic and phenotypic coefficients of variation were reported as the major tools used to
measure the variability that exists in a given population (Burton and Devane, 1988). The
larger environmental variance than genotypic variance signifies, the contribution of
environment was greater in the phenotypic expression of that trait. Abdel-aziz et al. (2015)
reported higher genotypic coefficient of variation with thousand seed weight and moderate
with number of seeds per plant in faba bean. They also reported low genotypic coefficient of
variation for number of pods per plant, number of seeds per pod and seed yield per plant.
Tafere Mulualem et al. (2013) also found high GCV for biomass and number of pods per
plant in their study of genetic variability, heritability and correlation in some faba bean
genotypes grown in Northwestern Ethiopia. Alghamdi (2007) also reported high genotypic
coefficient of variation for days to flowering, number of pods per plant and thousand seed
weight. Abdel-aziz et al. (2015) and Tafere Mulualem et al. (2013) reported higher
phenotypic coefficient of variation with number of pods per plant, thousand seed weight and
moderate for number of seeds per pod and seed yield per plant. High phenotypic coefficient of
variation was also reported for days to flowering, number of pods per plant and thousand seed
weight by Alghamdi (2007).

2.9. Heritability

Heritability can be defined, in broad sense, as the proportion of the genotypic variability to the
total variance (Allard, 1960). It refers to the portion of phenotypically expressed variation,
within a given environment and it measures the degree to which a trait can be modified by
selection.

According to Falconer and Mackay (1996) heritability in narrow sense is defined as “the ratio
of additive genetic variance to phenotypic variance”. Since broad sense heritability does not
give a clear picture of transmissibility of variation from generation to generation (because the
genetic variation includes the fixable and non-fixable dominance and epistatic variation), its
utilization is limited in plant improvement program. In contrast, estimate of heritability in a
narrow sense can give clearer picture than that of broad sense (Falconer and Mackay, 1996).
Estimation of heritability as a ratio of genotypic to phenotypic variance may vary greatly
depending upon the unit for which variance is considered (Johnson et al., 1955). The greater
the proportion of the total variability (i.e., due to environment), the more difficult will it be to
select for inherited differences. Conversely, if environmental variability is small in relation to
genotypic differences, selection will be efficient because the selected character will be
transmitted to its progeny. Generally, heritability indicates the effectiveness with which
selection of genotypes can be based on phenotypic performance (Johnson et al., 1955).

Singh (1990) categorized heritability into very high heritability (>80%), moderate high
heritability (50-80%), intermediate heritability (40-50%) and low heritability (<40%).
Therefore, selection for characters with heritability more than 80% should be fairly efficient
than for low heritability (<40%). This indicate that small environmental variability in relation
to genotypic differences, because, there is a close correspondence between genotype and
phenotype and relatively smaller contribution of environment to phenotype. For traits with
low heritability, selection may be difficult or virtually impractical due to the masking effect of
the environment on genotypic effects. Poehlmon and David (1995) described that, the
influence of the environment on the trait with high heritability is very low; and the increasing
value of broad sense heritability could serve as an indicator of selection based on phenotype,
particularly when it is accompanied by a relatively high variability and genetic advance.

Sufficiently very high heritability values were reported by Sharifi (2015) for traits like pod
length and pod width in genetic variation for seed yield and some of agro-morphological traits
in faba bean (Vicia faba L.) genotypes. Alghamdi (2007), Mellion Fikreselassie et al. (2012),
Tafere Mulualem et al. (2013) and Abdel-aziz et al. (2015) also reported moderately high
heritability values for traits such as plant height, number of pod per plant, number of seed per
plant, number of seed per pod, biomass weight per plot, thousand seed weight and seed yield.
Abdel Aziz et al. (2015) found moderate heritability for seed yield per plant and thousand
seed weight of faba bean genotypes and they also reported low heritability for traits: number
of pods per plant, number of seeds per plant and number of seeds per pod. Alghamdi (2007)
obtained the highest estimates of broad sense heritability for days to flowering, number of
pods per plant, number of seeds per plant and days to maturity.

2.10. Genetic Advance

Improvement in the mean genetic value of the selected plants over the base population is
usually termed as genetic advance under selection. It measures the difference between
genotypic values of generation obtained from the selected population over the mean value of
the population. Genetic advance under selection is a genotypic value which depends on three
things (Allard, 1960). These are genetic variability, heritability or masking effect of non-
genetic variability on the genetic variability and the selection intensity applied. Genetic
advance with conjunction to heritability can provide the estimate of expected gain for a
particular character (Johnson et al., 1955). The product of heritability, phenotypic standard
deviation and selection differential can estimate it. Burton and Devane (1953) indicated that
the genotypic coefficient of variation together with heritability estimate also gives the best
picture of expected advances from selection.

According to Allard (1960), the improvement in the mean genotypic value of the selected
plants over base population measures the expected genetic gain from selecting the best
performing genotypes for a given character. Johnson et al. (1955) reported broad sense
heritability along with genetic advances are usually more useful than heritability alone in
predicting the resultant effect of selecting the best individuals. Bakhiet et al. (2015) reported
high genetic advance with high heritability for traits: days to flowering, days to maturity,
plant height, number of branches per plant, number of pods per plant, number of seeds per
plant and seed yield per plot. Gemechu Keneni and Musa Jarso (2002) reported high genetic
advance as percent of mean along with high broad sense heritability for thousand grain weight
and seed yield per plot in comparison of faba bean genotypes. Kalia and Sood (2004) found
high heritability and high genetic advance for number of pods per plant.

2.11. Correlation of Traits

In plant breeding and genetic studies, correlated traits are of prime importance because of
linkage between genes or pleiotropic action of genes and changes brought by a natural or
artificial selection (Singh, 1993; Falconer and Mackay, 1996; Sharma, 1998). Correlated traits
are important for three basic reasons. First, in connection with the genetic causes of
correlation through the pleiotropic action of genes; second, in connection with the changes
brought about by selection; and third, in connection with the effect of natural selection on the
relationship of metric trait with its fitness, which is the primary agent, that determines the
genetic properties of that trait in a natural population (Falconer and Makey, 1996).

Inadequate knowledge of interrelationships among various traits and the practice of unilateral
selection for agronomic traits frequently end up with less than optimum result in plant
breeding (Bhatt, 1973). The practical utility of selecting for a given trait as a means of
improving another depends on the extent to which improvement in major traits is facilitated
by selection for the indicators. Such improvement depends not only on the genotypic
correlation but also phenotypic correlation (Johnson et al., 1955).Trait correlation studies
provide reliable information on the nature, extent and directions of selection (Kumar and
Chauhan, 1979). The knowledge of genetic correlations between different yield attributes
becomes of paramount importance when the breeder is confronted with problem of
introducing a quantitatively inherited trait into some agronomically superior cultivars from
wild or uneconomic genotypes (Saraf and Hedge, 1984). Seed yield is a polygenically
controlled complex trait and is dependent on a number of component traits that are also
quantitatively inherited. Selection on seed yield per se is often less effective, making it
imperative to go for indirect selection through component traits (Singh, 1983).
Generally, three types of correlations are involved between quantitative traits; and these are
phenotypic, genotypic and environmental correlations. The correlation between two traits that
can be directly observed is the correlation of phenotypic values or phenotypic correlations
(rp). Phenotypic correlations measure the extent to which the two observed traits are linearly
related. It is determined from measurements of the two traits in a number of individuals of the
populations. Genetic correlation (rg) is the associations of breeding values (i.e., additive
genetic variance) of the two traits. Genetic correlation measures the extent to which degree
the same genes or closely linked genes cause co-variation (simultaneous variations) in two
different traits. The correlation of environmental deviations together with non-additive genetic
deviations (i.e., dominance and epistatic genetic deviations) is referred to as environmental
correlations (re) (Singh and Chaudhary, 1977; Falconer and Mackay, 1996; Sharma, 1998).

Studies on genotypic and phenotypic correlations among traits of crop plants are useful in
planning, evaluating and setting selection criteria for the desired traits in breeding program
(Johanson et al., 1955). Correlations between different traits of crop plants may arise either
from genotypic or environmental factors. Environmental correlations arise from the effect of
overall environmental factors that vary at different environments. Correlations due to genetic
causes are mainly pleiotropic effects of genes and linkage (a phenomenon of genes inherited
together) between genes affecting different traits. Pleiotropy is the property of a gene, which
affects two or more traits; as a result it causes simultaneous variations in the two traits when
the genes are segregating (Singh, 1993; Falconer and Mackay, 1996).

Azarpou et al. (2012), Tafere Mulualem et al. (2013), Sharifi (2014) and Abdalla et al. (2015)
reported positive and significant phenotypic correlation between seed yield and thousand seed
weight. Tadele Tadesse et al. (2011) also found number of pods per plant, number of seeds
per pod, and thousand seeds weight had positive and significant phenotypic correlation with
seed yield per plot. In addition, Gemechu Keneni and Musa Jarso (2003) observed positive
and significant phenotypic correlation between seed yield and number of pods per plant.
Similarly, Alghamdi (2007) revealed positive and highly significant phenotypic correlation
between number of pod per plant and number of seed per plant and seed yield per plot.
Additionally, Azarpour et al. (2012), Sharifi (2014) and Abdalla et al. (2015) reported
positive and significant phenotypic correlation between number of pods per plant and traits
like plant height, seed yield per plot, number of seed per plant and thousand seed weight.
Moreover, Abdelmula et al. (2007) found positive and highly significant correlation between
harvest index and seed yield per plot. The report from different authors suggested that days to
flowering had positive and highly significant phenotypic correlation with plant height and
seed yield (Alghamdi, 2007; Bakhiet et al., 2015 and Sharifi, 2015). Similar to the previous
reports, Tafere Mulualem et al. (2013) found positive and highly significant phenotypic
correlation between days to flowering and days to maturity. Additionally, Tafere Mulualem et
al. (2013) and Alghamdi (2007) reported positive and significant phenotypic correlation
between days to maturity and number of branches per main stem and thousand seed weight.

2.12. Path Coefficient Analysis

Path coefficient analysis is simply a standardized partial regression coefficient which splits
the correlation coefficient into direct and indirect effects. In other words, it measures the
direct and indirect contribution of various independent traits on a dependent trait. Hence,
it measures the influence of a trait up on another trait; and permits the separation of the
correlation coefficient in to components of direct and indirect effects (Dewey and Lu, 1959).
With more traits in correlation studies, indirect associations become more complex and
important; consequently, correlation study coupled with path analysis will help to identify
yield attributing traits. Yield is a complex trait; and is influenced by its various components
directly as well as indirectly via other traits. Path coefficient analysis has indicated a direct
positive effect on seed yield by plant height days to flowering, time to maturity, branch
number, pod number, pod length, seeds per pod and seed weight. However, plant height, pod
number and seed weight had a direct negative effect on seed yield.

According to Shafique et al. (2016) the traits like pods per plant, plant height, days to
maturity, total weight of plant, primary branches per plant, seeds per plant, harvest index and
100-seed weight revealed positive direct influence on seed yield, whereas days to 50%
flowering showed negative direct effect on seed yield per plant. Chand et al. (2012) also
reported that harvest index and biological yield had high positive direct effect on seed yield.
Furthermore, it was observed that the highest indirect effects on seed yield were exhibited via
number of pods. In addition, Paneliya et al. (2017) path coefficient analysis revealed that
biological yield per plant and harvest index exhibited very high to high direct effect on seed
yield along with high indirect effect of number of pods per plant via biological yield per plant
and harvest index.

Additionally, Agrawal et al. (2018) revealed that biological yield, pods per plant, harvest
index, primary branches per plant and secondary branches per plant had high positive direct
effect on grain yield per plant under normal condition. Moreover, Bhanu et al. (2017) reported
negative direct effects of number of primary branches and number of seeds per pod on seed
yield. Whereas pod per plant, number of secondary branches, harvest index, plant height, days
to flowering, days to maturity and hundred seed weight had positive direct effects on seed
yield.

2.13. Cluster Analysis

Genetic divergence refers to genetic distance among the genotypes under consideration. It is
determined by using cluster analysis. D-square statistics (D 2) is one of statistical technique
developed by Mahalanobis (1936) used to classify the different genotypes into different
groups. The extent of diversity present between genotypes determines the extent of
improvement gained through selection and hybridization. The more divergent the two
genotypes suggests the more will be the probability of improving genotypes through selection
and hybridization.

According to Million Fikreselassie and Habtamu Seboka (2012) who used twenty-five elite
faba bean genotypes to study genetic variability of seed yield and yield related traits and
found high D2 value. Gemechu Keneni et al. (2007) also reported divergence between
Ethiopian germplasms. Birhanu Amare (2016) investigated genetic divergence of 32
genotypes of faba bean using Mahalanobis’s D 2 statistic and were grouped the genotypes into
seven clusters. In addition, Kumar et al. (2016) classified 65 genotypes in to nine different
non-overlapping clustering based on evaluations of 10 quantitative traits using Mahalanobis’s
D2 statistic. The number of clusters and the inter-cluster distances indicated presence of
considerably high genetic diversity in the genotypes under study. Behailu Mulugeta et al.
(2018) also reported thirty-two Ethiopian faba bean genotypes were grouped into five clusters
based on 23 agro-morphological traits. He found that the maximum distance was found
between C1 and C5 (D2 =554.60), while a minimum distance (D2 = 54.80) was observed
between C1 and C3. The high values of inter cluster distances indicated divergence among the
varieties and might be used in breeding programs for better genetic recombination and
selection of genetically divergent parents for exploitation in crossing programs. Similarly, Lal
et al. (2019) reported that 83 faba bean genotypes were grouped into ten different clusters by
using 12 agronomic traits and get high D2 value.

2.14. Principal Component Analysis

Principal component analysis (PCA) is a multivariate analysis that transforms a number of

possibly correlated variables into a smaller number of uncorrelated variables. It is a standard
tool in modern data analysis used to extract relevant information from confusing data sets by
identifying directions, called principal components (Chatfield and Collin, 1980). Therefore,
PCA used to identify and minimize the number of traits for effective selection and
improvement of yield and its related trait. The first principal component accounts more of the
variability in the data as possible followed by each succeeding component accounts for the
remaining variability as possible.

Birhanu Amare (2016) reported that the first four components accounted for 83.7% of the
total variation, whereas, the remaining 4 components accounted for only 16.3% of the
morphoagronomic diversity. PC I explained the most variability (36.1%), followed by PC II
(22.3%), PC III (15.4%) and PC IV (10.0%). In the first principal component, Days to
maturity, thousand seed weight, Number of seeds per pod and Number of Tillers per Plant
were more important traits contributing more to the variation. In his experiment, the PC
analysis ultimately showed the amount of variability for the traits that could be used for the
improvement of faba bean genotypes. Rebaa et al. (2017) performed a PCA among 21
populations of faba bean prevailing in Tunician region of Australia for a number of
quantitative traits including, days to flowering, plant height (cm), number of stems per plant,
pods number per node, seeds number per pod, pod length (cm), 100-seed weight (g). They
found that, the first three principal components (PCs) accounted for 40.56% of the total
variation, of which PC1, PC2 and PC3 explained 20.64, 11.22 and 8.70% of the variation,
respectively. Behailu Mulugeta et al. (2018) also reported that the principal components
analysis (PCAs) with Eigen value greater than one contributed 85% of the entire diversity
among the genotypes. The first three PCs contributed 66% (PC1 = 38%, PC2 = 19% and PC3
= 9%) of total variation among Ethiopian faba bean varieties.

Tiwari and Singh (2019) who reported the first three principal components exhibited eigen
values more than one and explained about 75.53% of the total variations present in the data,
so first three PCs were given due importance for further explanation. First principal
component considered as the most important component which explained 44.00% of the total
variance. Important eigenvectors for PC1 were days to maturity, plant population per plot, No.
of branches per plant, No. of pods per plant, pod width and No. of seeds per pod. The second
principal component contributed 15.60% of the variation among genotypes. PC2 was
positively defined by days to maturity, while negative influence was noticed in 100 seed
weight. The third principal component accounted for 15.13% of the total variance, and was
positively influenced by plant height and pod length.
 Chapter 3. MATERIALS AND METHODS

3.1. Description of the Study Area

The experiment was conducted at Kudmi, Bahir Dar University research site in Mecha district
of West Gojjam zone of Amhara regional state, Ethiopia in 2019 main cropping season.
Mecha is located about 525 km North East of Addis Ababa and 34 km South West of Bahir
Dar, the capital city of Amhara region. It is located at latitude of 10°30' N and longitude of
37°29' E. The climatic condition alternates between summer rainfall and winter dry season
with mean annual rainfall of 1572 mm. The mean temperature ranges between 24°C and 27
°C; and the altitude is 2009 m.a.s.l. Agriculture is the main economic sector in the study area.
The main agricultural activities currently practiced include irrigation (modern and traditional)
and mixed farming. The major crops cultivated around the study area are maize, teff, wheat
and legumes (Source; Mecha Woreda agricultural office).

3.2. Experimental Materials and Design

Totally, 81 faba bean genotypes were used in the present study. Seventy-eight of the
genotypes were accessions obtained from Ethiopian Biodiversity Institute; the two were
standard checks obtained from Adet Agricultural Research Center; and one was a local check
from local source (Appendix Table 1). The design used was 9 X 9 simple lattice. Each
accession was planted on two rows. Spacing between rows and within rows were 40 cm and
10 cm, respectively. The single plot size was 0.8m x 1m (0.8m 2). Spacing between blocks was
1 meter.

3.3. Data Collected

The pre- and post-harvest observations was recorded from randomly taken five plants from
each plot in each replication for all the traits considered except for days to flowering, days to
maturity, biomass yield, grain yield, harvest index which were determined based on plot basis.
All agronomic traits of sampled plants and seeds were taken at the right growth stage of the
plant for each trait.

3.3.1. Data collected on plant basis

Plant height (cm): was measured from five randomly taken plants from the ground surface to
the tip of the main stem and the average was used for further analysis.
Pod length (cm): was measured at harvesting from five randomly taken pods of the five
randomly taken plants and the average was used for further analysis.
Number of pods per plant: was recorded from all pods borne on the five randomly taken
plants and averaged.
Number of seeds per pod: was recorded from seeds of five randomly taken pods from the
five randomly taken plants and averaged to represent the plot.
Number of branches: was counted and averaged from branches extending from the main
stem of the five randomly taken plants.
Disease score: incidence of disease was scored using 1-9 scale method; where, 1 refers highly
resistance and 9 indicate highly susceptible.

The response of tested accessions were classified into six reaction groups according to Abo-
Hegazy et al. (2012), where 0 to 2% is highly resistant (HR), > 2 to 15% is resistant (R), > 15
to 40% is moderately resistant (MR), > 40 to 60% is moderately susceptible (MS) and > 60 to
80% is susceptible (S) >80 to 100 % is highly susceptible (HS) based on percent disease
severity values. The disease data recorded based on scoring scale was converted to percentage
severity index (PSI) according to Wheeler (1969). Percentage Severity Index (PSI) used to
convert scale data into percent and ease of analysis. Percent severity index (PSI %) was
calculated using:

∑( NPC∗CR)∗100
PSI (%) =
NIP∗MSC

Where; NPC = number of plants in each class rate,

CR = class rate,
NIP = number of infected plants and
 MSC = maximum severity class rate.
Table 3.1. Rating scale of diseases
Chocolate spot scale
1=when no disease symptoms or very small specks (highly resistance) was seen
3=few small disease lesions (resistant) was seen
5 =some coalesced lesions, with some defoliation (moderately resistant) was seen
7=Large coalesced sporulating lesions, 50% defoliation and some dead plants (susceptible)
9=Extensive, heavy sporulation, stem girdling, blackening and death of more than 80% of
plants (highly susceptible)
Rust scale
1=No pustules or very small non-sporulating flecks (highly resistant)
3 =Few scattered pustules covering less than 1% of the leaf area, and few or no pustules on
stem (resistant)
5 =Pustules common on leaves covering 1-4% of leaf area, little defoliation and some
pustules on stem (moderately resistant).
7 =Pustules very common on leaves covering 4-8% of leaf area, some defoliation and many
pustules on stem (susceptible).
9 =Extensive pustules on leave, petioles and stem covering 8-10% of leaf area, many dead
leaves and several defoliations (highly susceptible).
Ascochyta blight scale
1=when no disease symptoms or very small specks (highly resistance) was seen
3=few small disease lesions (resistant) was seen
5 =some coalesced lesions, with some defoliation (moderately resistant) was seen
7=Large coalesced sporulating lesions, 50% defoliation and some dead plants (susceptible)
9=Extensive, heavy sporulation, stem girdling, blackening and death of more than 80% of
plants (highly susceptible)
Source: Bernier et al. (1993).
3.3.2. Data collected on plot basis

Days to 50% flowering: was recorded from date of sowing to the date of which 50 percent of
the plants on a plot opened their flowers.
Days to 75% maturity: was recorded from date of sowing to the stage when 75 % of the
plants in a plot changed the color of their pods from green to yellow.
Grain yield (g/plot): obtained from each plot was used to estimate grain yield per hectare
after cleaned and adjusted to10% moisture content.
100-seed weight (g/plot): 100-seeds was taken randomly on each plot and weighed to
represent this trait.
Biological yield (g/plot): was recorded from the above ground parts of harvested plants which
was sun dried and weighed to get the biological yield per plot in grams.
Harvest Index: was calculated by dividing grain yield by the biological yield. Harvest Index
grain yield ( g )
(HI) = *100
biological yield ( g )

Qualitative traits
Seed size (boldness) - were classified into three categories small, medium and bold (1= Small,
2 = Medium and 3 = Bold).
Seed shape - was classified into three categories, oblong, semi-spherical and semi-ovals (1 =
Oblong, 2 = Semi spherical and 3 = Semi oval) to know the degree of diversity in this trait
Seed coat colour - was also classified into light green, pale green and light brown to copper
colour (1 = Light green, 2 = Pale green and 3 = Light brown to copper colour)
Then, such qualitative data were converted into percentage through;
No. of accessions ∈a scale
∗100
Total no . of tested accessions

3.4. Data Analyses

The data for each trait was subjected to analysis of variance (ANOVA) for simple lattice
design. Analysis of variance was done using the procedures outlined by Gomez and Gomez
(1984) with the help of SAS software version 9.2 (SAS 2008). Analysis of variance was done
using lattice (Proc lattice) and general linear models (Proc GLM) procedures of SAS
software. General linear models were used to calculate adjusted block sum of squares,
adjusted treatment sum of squares and intra block error; and handle relating one or several
continuous dependent variables to one or several independent variables. PROC GLM was
analyzed within the framework of general linear models. Proc lattice was used to analyze
experiments from lattice design and produce more accurate estimates of the treatment effects.
The relevant values were generated on plot and plant bases. Duncan’s Multiple Range Test
(DMRT) was used for means separation.

Model of the Simple Lattice Design is as follows:

Yijl = μ + Ti + yj + pl(j) + Σijl

Where; Yijl = 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 (mean),
Ti = treatment effect i=1, 2, 3 ...t,
yj = replication effect j= 1, 2 ...r,
pl (j) = block within replicate effect l= 1, 2...b and
Σijl = the error associated to the observation Yil (j)
Table 3.2. The structure of analysis of variance (ANOVA) for simple lattice design
Source of variation DF SS MS Computed F

Replication r-1 SSR MSR MSR/MSE

Treatment (unadj.) K2-1 SST (unadj.) MST(unadj.) MST/MSE
Treatment (adj.) K2-1 SST(adj.) MST(adj.) MST/MSE
Block within replication(adj.) r(k-1) SSB(adj.) MSB(adj.) MSB/MSE
Intra–block error (k-1)(rk-k-1) SSE MSE
Total rk2-1 SST

Note: DF = degree of freedom, SS = sum square, MS = mean square, r = number of

replications, k2= number of treatments and k = number of plots in a block
Source: Gomez and Gomez (1984)

3.4.1. Estimation of variance components

The phenotypic and genotypic variance of each trait was estimated from the analysis of
variance. The expected mean squares under the assumption of random effects model was
computed from linear combinations of mean squares. Random model was used when dealing
with random sample from some population to make inference. Range, mean, standard error,
phenotypic and genotypic variance and coefficient of variation were calculated to estimate
variability. Genotypic variance (σ2g), phenotypic variance (σ2p), phenotypic coefficient of
variation (PCV) and genotypic coefficient of variation (GCV) were estimated using the
formula adopted from Burton and Devane (1953):

MSg−MS e
Genotypic variance (σ2g) =
r

Environmental variance (σ2e) = MSe

Where; MSg and MSe are the mean square of genotypes and error from the analysis of
variance, respectively. r is the number of replications. Then, the phenotypic variance was
estimated as the sum of the genotypic and environmental variance.

Phenotypic variance (σ2p) = σ2g+ σ2e

Coefficients of variation at phenotypic and genotypic levels were estimated using the
following formula:
Genotypic Coefficient of Variation (GCV) = (√σ2g /grand mean) x 100
Phenotypic Coefficient of Variation (PCV) = (√σ2p/grand mean) x 100

Heritability in the broad sense

Broad sense heritability (H2) is the ratio of the genotypic variance (σ 2g) to the phenotypic
variance (σ²p) and was computed on genotype mean basis according to Allard (1960) and
Fisher (1987) as follows:

𝐻² = 𝑥100
σ²g
σ² p
Where; σ2g = (MSg - MSe) /r
σ²p = σ²g +σ²e
σ²e = Mse and MSg- and MSe are the mean squares of genotypes and error, respectively.

As Johnson (1955) stated the ranges of heritability were categorized as; low = 0 - 30, medium
= 31 - 60 and high > 61
Genetic advance

The methods illustrated by Johnson (1955) were used to compute expected genetic advance
(GA) and GA as percent of the mean assuming selection of the superior 5% of the genotypes.

GA=K *𝜎𝑝*h2 and GA as % of the mean = (GA/X) ×100

Where; h2 = the heritability ratio,
X = grand mean of the trait being evaluated,
𝜎𝑝 = phenotypic standard deviation and
k = constant which varies on the selection intensity.

3.4.2. Correlation analysis

Phenotypic correlation, the observable correlation between variables, which is the sum of
genotypic and environmental effects were calculated from variance covariance components
using the formula of Miller et al. (1958) as follows:

Genotypic correlation= (Genotypic covariance of X and Y) / √ (σ2g X σ2g Y)

Phenotypic correlation= (Phenotypic covariance of X and Y) / √ (σ2 p X σ2pY)
Where,σ2p X = Phenotypic variance for character X
σ2p Y = Phenotypic variance for character Y,
σ2g X = Genotypic variance for character X, and
σ2g Y = Genotypic variance for character Y

The significance of phenotypic correlation coefficients was tested by the formula of Singh and
Chaudhary (1985).
t’= rpxy √ (n-2 / 1-rp2xy)

t’ value was tested against the tabulated t-value for (n- 2) degree of freedom where n was the
number of genotypes studied; whereas the coefficient of variation at genotypic level was
tested for significance using the formula proposed by Robertson (1959).
r g xy
t=
SE r g xy
The calculated‘t’ value was compared with ‘t’ tabulated at (n-2) degree of freedom at 1% and
5% levels of significances.
Where;
Where; SEr xy is the standard error of genotypic correlation coefficient; Hx and Hy are
g

heritability for traits x and y, respectively.

3.4.3. Path coefficient analysis

Path coefficient analysis is a tool to partition the observed correlation coefficient into direct
and indirect effects of yield components on grain yield. Path analysis provides precise picture
of character associations for formulating efficient selection strategy. Path coefficient analysis
differs from simple correlation that it points out the causes and their relative importance,
whereas, the later measures simply the mutual association ignoring the causation. The concept
of path coefficient was developed by Wright, (1921) and the technique was first used for plant
selection by Dewey and Lu, (1959). Path analysis has emerged as a powerful and widely used
technique for understanding the direct and indirect contributions of different characters to
economic yield in crop plants.

Based on genotypic correlation, path coefficient which refers to the direct and indirect effects
of the yield attributing traits (independent trait) on grain yield (dependent trait) were
calculated as the method used by Dewey and Lu (1959) as follows:
rij = Pij + Σ rikpkj
Where; rij = mutual association between the independent character (i) and dependent
character (j) as measured by the genotypic correlation coefficients.
Pij = direct effects of the independent character (i) on the dependent variable (j) as
measured by the genotypic path coefficients, and
Σrikpkj = Summation of components of indirect effects of a given independent
character (i) on a given dependent character (j) via all other independent
characters (k).
Residual factor (R), the contribution of the remaining unknown factor was estimated using
Singh and Chaudhury (1985) method:
R = √ (1− ΣPij rij)
3.4.4. Clustering of accessions and genetic divergence analysis

Cluster analysis was made using SAS Software version 9.2 (SAS Institute, 2008). Based on
the squared distances (D2) values, clustering of accessions was done using Tocher’s method as
described by Singh and Chaudhary (1985). Unweighted Pair Group Methods with Arithmetic-
average (UPGMA) based on the generalized D 2 distances by average linkage method of
agglomerative hierarchical clustering (the two closest clusters are merged into a single new
cluster) built to group the faba bean genotypes into genetically distinct classes based on
phenotypic traits. Appropriate number of clusters was determined by using points where local
peaks of Pseudo F-statistics join with small values of the Pseudo t 2 statistics followed by a
larger Pseudo t2 for the next cluster fusion. The dendrogram was constructed on the basis of
average linkage and squared distance used as a measure of dissimilarity technique using JMP
software.

Genetic distance between pair of clusters was calculated using the generalized Mahalanobis's
D2 statistics based on the recommendation of Singh and Chaudhary (1985). Average intra
2
D i
cluster D2 values were estimated using the formula where, Di2 is the sum of distance
n
between all possible combinations (n) of the accessions included in a cluster. The D 2 statistics
measures the force of differentiation at intra cluster and intercluster levels (Singh, 2007).
Hence, genetic distance between clusters as standardized Mahalanobis’s D 2 statistics
(Mahalanobis, 1936) were calculated as follow:
D2ij = (Xi- Xj) S-1 (Xi – Xj)

Where; D2ij = the square distance between any two genotypes i and j,
Xi and Xj = the vectors for the values for genotype ith and jth genotypes, and
S-1 = the inverse of pooled variance covariance matrix.

The significance level of genetic distance between and within clusters was tested at both 1%
and 5% level of probability using Chi-square test. The 𝐷2 values obtained for pairs of cluster
using SAS statistical package were considered as the calculated values of Chi-square (χ 2) and
tested for P degree of freedom, where P is the number of traits considered (Singh and
Chaudhary, 1985).

3.4.5. Principal component analysis

Principal component analysis (PCA) was used to find out the traits, which accounted more to
the total variation. The data were standardized to mean zero and variance of one before
computing principal component analysis. Principal components were calculated based on
correlation matrix using SAS computer software. As suggested by Johnson and Wichern
(1988), eigenvector greater than half divided by the standard deviation (square root) of the
eigen value of the respective PC was employed as general guideline for weighing the relative
significance of traits constituting to the PCs.
 Chapter 4. RESULTS AND DISCUSSION

4.1. Variability among Accessions

Mean squares of the 14 traits from analysis of variance (ANOVA) are presented in Table 4.1.
Highly significant (p<0.01) differences among accessions were observed for days to
flowering, plant height, pod length, number of pods per plant, number of branches per plant,
biomass yield, grain yield, 100 seed weight, harvest index, ascochyta blight, chocolate spot
and rust disease score. This highly significant difference indicates the existence of variability
among accessions for traits studied. The presence of variability in the accessions offers an
opportunity in improvement of yield and its contributing traits through selection in Faba bean.
Similar results were obtained by Gemechu Keneni et al. (2005) in 160 faba bean accessions.
Highly significant variations for all the yield related traits were also reported by Alghamdi
(2007). Hamza (2017) and Abdullah (2018) also reported significant variability among the
faba bean genotypes. The analysis of variance revealed that, the relative efficiency of lattice
design over RCBD was not efficient for all traits except plant height and hence implies RCBD
was good than lattice design.
.
 Table 4.1. Analysis of variance for 14 traits of eighty-one faba bean accessions

Traits Mean square CV (%) R2 R.E. to RCBD

Rep (DF=1) Accessions (DF=80) Rep x Block (16) Error (DF= 64)
DF 0.10ns 18.57** 3.02ns 2.21 3.30 0.92 101.87
DM 43.56ns 25.29ns 28.53ns 18.28 3.64 0.68 103.76
PH 365.70ns 230.18** 212.36** 109.68 10.75 0.77 108.26
PL 2.57** 0.53** 0.24ns 0.19 11.74 0.80 101.12
NPP 1.03ns 45.44** 7.25ns 6.74 22.36 0.89 100.10
NSP 1.81** 0.17ns 0.09ns 0.12 12.32 0.68 95.91
NBP 0.00ns 0.61** 0.26* 0.17 24.16 0.81 103.24
BY 24966.00ns 9567533.60** 817978.00ns 691793.00 10.23 0.95 100.55
GY 31441.00ns 810072.84** 10553.00ns 14290.00 5.72 0.99 94.77
ns
HSW 6.97 173.27** 47.64* 28.96 12.93 0.89 104.69
ns ns
HI 3.21 106.81** 16.36 12.71 14.44 0.90 101.23
ns ns
AB 12.20 532.02** 0.04 6.19 7.07 0.99 97.23
ns ns
CS 3.05 547.10** 0.33 24.49 15.07 0.96 104.92
RT 76.22* 590.28** 0.12ns 14.48 14.30 0.98 100.00
NOTE: ** = Highly significant (p<0.01), * = Significant (p<0.05), ns = Non-significant, CV = Coefficient of variation, DF= days to flowering, DM= days to
maturity, PH= plant height(cm), PL= pod length, NPP= number of pods per plant, NSP=number of seeds per pod, NBP= number of branches per plant, BY=
biomass yield, GY= grain yield, HSW= 100 seed weight, HI= harvest index, AB= ascochyta blight, CS= chocolate spot, RT= rust disease, R.E to RCBD=
relative efficiency to RCBD.

32
4.2. Estimation of Mean and Range

The range and mean performance of accessions for the 14 traits are shown in Table 4.2. The
grain yield ranged from 1148 to 5167 kg ha -1 with a mean value of 2050 kg ha -1. The highest
grain yield was recorded in accession 2354333 (5167 kg/ ha-1) while the lowest grain yield
was obtained from accession 208085 (1148 kg/ ha-1). This indicates the existence of a wide
range of variability among accessions in the trait of grain yield. Therefore, accessions 235433,
235709, 219089, 25325, 25310, and 25290 were promising for the grain yield improvement of
faba bean in breeding program. The results of present study are in conformity with earlier
findings of Tamene Temesgen (2015) who reported the highest yield of 6649 kg ha -1 was
obtained from variety ‘Degaga’ at Kulumsa in 2009, while the lowest was 1198 kg ha -1 from
the varieties ‘Kuse-2-27-33’ at Bekoji in 2007. Behailu Mulugeta (2016) who reported the
minimum and maximum average seed yield ranged from 2955.59 kg ha -1 to 5631.46 kg ha-1.
Teame Gereziher et al. (2017) also reported that the highest grain yield was obtained on
varieties Dosha and Tumsa, which was 3891 and 3437 kg ha -1, respectively. Ashenafi Mitiku
and Wolde Mekuria (2015) had also reported yield variation from 3703.7-4886.8 kg ha -1 and
3436.2-4701.6 kg ha-1 in Agarfa and Sinana trial sites, respectively.

For the phenological trait, days to flowering ranged from 42 to 59 with a mean value of 45.56
days. The results indicate the presence of high variability among the accessions for days to
flowering. Accordingly, accession 25018 was the late flowering whereas the local was earliest
in flowering. This result agreed with Tafere Mulualem et al. (2012) who reported that days to
flowering had significantly affected by variety. Days to maturity ranged from 109 to 131 with
a mean value of 118.20 days. Accession 240497 was earliest maturing and Dosha was the late
maturing. Biological yield is an indicator of ability of plant to adopt well under certain agro-
climatic conditions. It is partially, true because sometime plant failed to produce seed and
unable to complete its life cycle successfully. The tested accessions have got good amount of
variability in respect to biological yield. Biological yield shows a range of 2471 to 15278 kg
ha-1 with a mean of 8310 kg ha-1. The maximum biological yield was recorded from Dosha
and minimum from accession 212576. This result indicates that there is ample scope for
improvement in this important trait. This trait indicates that the crop has good plasticity and

33
having great potential to improve its performance. These results are in the close conformity
with the findings of previous researchers (Bora et al., 1998; Veasey et al., 1999; Singh et al.,
2017). Plant height ranged from 53.6 to 132 cm with a mean of 97.75 cm. The maximum and
minimum plant heights were recorded from local and accession 212568, respectively. This
indicates the existence of variability among accessions. This result was in line with the work
of Teame Gereziher et al. (2017) who reported that Hachalu (120.70 cm) was the longest
plant height. The minimum and maximum number of branches per plant ranged from 0.6 to
3.8 in accession 25006 and 255709, respectively.

Number of pods per plant ranged from 3.5 to 39.2 with a mean of 11.70. The maximum
number of pods per plant was obtained from accession 235433 while the minimum was from
accession 212575. This trait clearly shows that there is presence of sufficient variability in the
accessions. Number of pods per plant can be important selection criterion for the development
of high yielding accessions as it is important yield component in faba bean. Tafere Mulualem
et al. (2012) and Teame Gereziher et al. (2017) reported the significant difference among faba
bean varieties with regard to the number of pods per plant. Likewise, Tewodros Tesfaye et al.
(2015) and Degife Asefa and Kiya Adare (2016) had reported significant differences in the
number of pods per plant, 100 seed weight, as well as grain yield among the faba bean
varieties at Gamo Gofa and North Gonder zones respectively. Number of seeds per pod
ranged from 1 to 3.8 with a mean value of 2.77. Maximum number of seeds per pod was
obtained from accession 25274 whereas accession 25018 produced minimum number of seed
per pod.

Hundred seed weight ranged from 24 to 101.2 gram per hundred seed; Dosha had the
maximum and accession215748 had the minimum 100 seed weight. Ashenafi Mitiku and
Wolde Mekuria (2015) also reported significant difference among faba bean varieties in 100
seed weight. Harvest index is an effective indicator of source to sink relation and translocation
of photosynthesis. The range of HI was 12.27 to 55.13% with mean value of 25.71. This
shows the range of HI with good amount of variability. The results of present study are in
conformity with earlier findings of (Bora et al., 1998; Veasey et al., 1999; Singh et al., 2017).
Pod length ranged from 2.3 to 6.5 cm with a mean value of 3.75 in accession 25006 and

34
Dosha, respectively. This indicates the presence of variability among accessions and selection
can result in the improvement of pod length of faba bean. Similar results were reported by
Singh et al. (2017).

Table 4.2. Minimum, mean and maximum values of fourteen traits of 81 faba bean accessions

Minimu Maximu
Traits Mean Std Dev m Acc. m Acc.
DF 45.56 3.54 42 Local 59 25018
DM 118.20 5.12 109 240497 131 Dosha
PH 97.75 14.64 53.6 212568 132 Local
PL 3.75 0.65 2.3 25006 6.5 Dosha
NPP 11.70 5.31 3.5 212575 39.2 235433
NSP 2.77 0.41 1 25018 3.8 25274
NBP 1.70 0.62 0.6 25006 3.8 235709
8310.0 2426.00 2471 212576 15278 Dosha
BY 0
2050.0 675.77 1148 208085 5167 235433
GY 0
HSW 41.86 11.06 24 215748 101.2 Dosha
HI 25.71 7.91 12.27 25006 55.13 27290
AB 35.53 16.84 11.11 235433 77.78 215748
CS 33.74 17.60 11.11 240497 77.78 215748
RT 26.34 18.11 11.11 25274 77.78 212576
Note: DF= days to flowering, DM= days to maturity, PH= plant height, PL= pod length, PP= number of pod per
plant, SP=number of seed per pod, NBP= number of branch per plant, BY= biomass yield, GY= grain yield,
HSW= 100 seed weight, HI= harvest index, AB= ascochyta blight, CS= chocolate spot, RT= rust disease,
Acc=accession.

Disease score for Ascochyta blight, chocolate spot and rust were ranged from 11.11 to
77.78% for each based on percentage severity index (PSI) conversion. Treatment 10, 13, 17,
19, 36, 38, 43, 49, 62, 63, 67, 68, 69, 72, 78, and 80 were resistant for Ascochyta blight
disease but treatment 7, 14 and 45 were susceptible by Ascochyta blight. The response of
tested treatment 10, 43, 49, 62, 63, 72, 78, 11, 24, 41, 46, 50, 52, 58, 64, 70, 71, 74, 4, 21 and
81 were resistance of chocolate spot however treatment 45, 7, 8, 54 and 32 were susceptible.

35
Most of accessions categorized under resistance and moderately resistant for rust disease but
treatment 18, 29 and 61 were susceptible. These shows the accessions were significantly
different for disease traits. This result is conformity with (Degife Asefa and Kiya Adare,
2018; Abo-Hegazy et al., 2012) who classified faba bean genotypes into different reaction
group. Similar results also reported for disease resistance traits by (Zakaria et al., 2015;
Eldemery et al., 2016 and Belal et al., 2018). The overall mean performances of accessions
were listed in Appendix Table 2.

Seed size (boldness of seed) in small scale containing 37 accessions or 45.68% among studied
accessions (Table 4.3.). Medium seed size had largest number of accessions which is 39
(48.15%) and bold seed size contains 5 number of accessions or 6.17% out of 81 tested
accessions. Considerable amount of variability existed in this trait. So this trait has a good
opportunity to improve the boldness of seeds through selection. Sheelamary et al. (2015) also
characterized 50 faba bean accessions using qualitative traits of seed coat colour and seed
shape. Seed shape in the first oblong seed shape was shown in 25 accessions, 30.86% of the
accessions. Semi-spherical and semi-ovals were seen in 35 (43.21%) and 21 (25.93%)
accessions, respectively.

Light green seed coat colour was observed in10 accessions. Majority of accessions (39) had
pale green seeds and 32 of accessions showed light brown class. Faba bean plants vary widely
in expression of yield and its contributing traits along with quality parameters that offers
opportunities in faba bean improvement, which were also reported by Kalia and Pathania
(2007) and Ammar et al. (2015). Generally, the qualitative traits used clearly showed the
presence of variability among accessions. These indicate that these traits can be used as
selection criterion for the improvement of faba bean accessions. Similar results were obtained
by Singh et al. (2017).

36
Table 4.3. Extent of variability in qualitative traits among faba bean accessions

Traits Scale Number of accessions Percentage of accessions (%)

Seed size 1 37 45.68
2 39 48.15
3 5 6.17
Seed shape 1 25 30.86
2 35 43.21
3 21 25.93
Seed coat colour 1 10 12.35
2 39 48.15
3 32 39.51

Note: Seed size (1= Small, 2 = Medium and 3 = Bold); Seed shape (1 = Oblong, 2 = Semi spherical
and 3 = Semi oval);Seed coat colour (1 = Light green,2 = Pale green,3 = Light brown to copper
colour).

scale 1 2 3

48.15% 48.15%
45.68%
43.21%
39.51%

30.86%
25.93%

12.35%
6.17%

Seed size Seed shape Seed coat colour

37
Figure 4.1. Diagrammatic representation of qualitative traits observed in 81 Faba bean
accessions.

4.3. Estimates of Genetic Parameters

4.3.1. Phenotypic and genotypic coefficients of variation

The estimates of genotypic (σ2g) and phenotypic (σ2p) variances and phenotypic (PCV) and
genotypic (GCV) coefficients of variation, heritability (broad sense), genetic advance and
genetic advance as percent of mean were determined for the traits considered, and are
presented in Table 4.4. According to Deshmukh et al. (1986), PCV and GCV values more
than 20% are considered as high, values between 10 and 20% as medium and values less than
10 % as low. Accordingly, high PCV was observed for 100 seed weight, biological yield,
harvest index, grain yield, number of branches per plant, number of pods per plant, ascochyta
blight, chocolate spot and rust disease score. These indicate the existence of wide phenotypic
variation among the accessions considered in the present study and showed the possibility of
genetic improvement through selection regarding those traits. Similar findings were reported
by Tafere Mulualem et al. (2013) for 100 seed weight, disease resistance, number of pods per
plant, grain yield and biological yield.

Medium PCV was observed for traits like plant height and pod length indicating the existence
of lesser phenotypic variability among faba bean accessions. Similar results were reported by
Kumar et al. (2017) that PCV was medium for traits like plant height and pod length. In the
present study, low PCV was observed for days to flowering which indicate the presence of
narrow genetic variation. However, the improvement of this trait could be possible through
hybridization or induced mutagenesis followed by selection. Similarly, Dev et al. (2017)
reported low PCV for days to flowering in chickpea.

High GCV was observed for traits like number of pods per plant, number of branches per
plant, biological yield, grain yield, 100 seed weight, harvest index, ascochyta blight, chocolate
spot and rust disease score. These results indicate the existence of considerable genotypic
variability among faba bean accessions for these traits and greater influence of genetic factors

38
 for the expression of these traits suggesting the possibility of improving these traits through
selection.

Table 4.4. Estimates of variances, coefficient of variation, heritability and genetic advance of
traits

2 2
Traits σ g σ p GCV (%) PCV (%) H2 (%) GA GAM (%)

DF 8.18 10.39 6.28 7.08 78.71 5.23 11.47

PH 60.25 169.93 7.94 13.34 35.46 9.52 9.74

PL 0.17 0.36 11.02 16.00 47.48 0.59 15.65

NPP 19.35 26.09 37.60 43.66 74.16 7.80 66.70

NBP 0.22 0.39 27.52 36.81 55.89 0.72 42.38

4437870.3
BY 0 5129663.30 25.35 27.25 86.51 4036.43 48.57

GY 397891.42 412181.42 30.77 31.32 96.53 1276.70 62.28

HSW 72.15 101.12 20.29 24.02 71.36 14.78 35.32

HI 47.05 59.76 26.67 30.06 78.72 12.54 48.76

AB 262.91 269.11 45.64 46.17 97.70 33.08 93.11

CS 261.31 285.79 47.90 50.10 91.43 32.98 97.73

RT 287.90 302.38 64.43 66.03 95.21 34.61 131.43

Note: DF= days to flowering, PH= plant height, PL= pod length, NPP= number of pods per plant, NBP= number
of branches per plant, BY= biomass yield, GY= grain yield, HSW= 100 seed weight, HI= harvest index, AB=
ascochyta blight, CS= chocolate spot, RT= rust disease, GV=genotypic variance, PV= phenotypic variance,
GCV= genotypic coefficient of variance, PCV= phenotypic coefficient of variance,H2= broad sense heritability,
GA= genetic advance and GAM (%)=genetic advance as percent of mean.

The result is in conformity with Kumar et al. (2017) who reported high GCV for number of
pods per plant, number of branches per plant, 100 seed weight. Medium GCV was observed
for pod length and low GCV revealed for traits like days to flowering and plant height. These
results indicate greater influence of environmental factors for the expression of these traits.

39
Kumar et al. (2017) reported medium GCV for pod length. The low GCV estimates observed
in the present investigation indicates the presence of narrow genetic variation on the traits.
The result agrees with the findings of Dev et al. (2017) who reported low GCV for plant
height in chickpea. Similar findings were reported by Saghin (2002) and Kalia (2003) for days
to flowering in faba bean. The amount of genotypic and phenotypic variability existing in a
species is important to improve traits by direct selection. The extent of the environmental
influence on any trait has been determined by the magnitude of the differences between the
genotypic and phenotypic coefficients of variation. Large differences reflect high
environmental influence, while small differences reveal high genetic influence. In this point
of view, results for all the studied traits, revealed a little higher phenotypic coefficient of
variation than genotypic coefficient of variations. Hence, slight differences might be indicated
minimum environmental influence and consequently greater role of genetic factors on the
expression of traits.

4.3.2. Broad sense heritability

Heritability (H2) in broad sense estimates were generally high for most studied traits which
ranged from 35.46 % for plant height to 97.70 % for ascochyta blight disease score (Table
4.4.). Accordingly, in the present study, high broad sense heritability estimates were found for
traits such as 100 seed weight, number of pods per plant, days to flowering, harvest index,
biomass yield, chocolate spot, rust, grain yield, and ascochyta blight disease score. The
results indicated that selection based on phenotypic expression of individual accessions for
these traits may be easy due to relatively small contribution of the environment to the
phenotype. The high H2 values of these traits indicate the high possibility of transferring traits
from parents to the next progeny. Those traits that showed high and medium heritability are
found to have high GCV values. Selection for these traits is relatively easy because most of
the variation is genetic rather than environmental. In agreement with these results, high
estimates of broad sense heritability was reported for 100-seed weight, grain yield and
chocolate spot disease in faba bean genotypes by earlier workers (El-Badawy et al.,2012;
Million Fikreselassie et al., 2012; Tafere Mulualem et al., 2013; Hamza et al., 2017).

40
Sheelamary and Shivani, (2015) also observed high heritability estimates for days to 50%
flowering.

In the present study medium heritability estimates were recorded for traits like plant height,
pod length and number of branches per plant. The traits with high to moderate heritability
may respond moderate to high for phenotypic selection (Singh et al., 1996). However,
selecting superior individuals based on heritability estimates alone may not be evidence for
genetic improvement. For traits with low heritability, selection may be difficult or virtually
impractical due to the masking effect of the environment on genotypic effects (Johnson et al.,
1955). Low heritability was also reported by earlier workers Kalia and Sood (2004),
Alghamdi (2007), Tafere Mulualem et al. (2013).

4.3.3. Genetic advance

Genetic advance as percent of mean varied from 9.74% for plant height to 131.43% for rust
disease score (Table 4.4.). High heritability was recorded for number of pods per plant,
biomass yield and 100 seed weight. The traits that exhibit high heritability coupled with high
genetic advance could be useful in the selection process as they are controlled by additive
genes and less influenced by environment (Panse and Sukhatme, 1985). The report by Kalia
and Sood, (2004) showed high heritability and high genetic advance for number of pods per
plant. Similar results were reported by Behailu Mulugeta et al. (2018) for number of pods per
plant, biomass weight per plot and thousand seeds weight.

According to Johnson et al. (1955) high heritability estimates along with the high genetic
advance as percent of mean is usually more helpful in predicting gain under selection than
heritability alone. Panse (1967) reported that high heritability associated with high genetic
advance was indicative of additive gene effects and high heritability with low genetic advance
was an indication of dominance and epistatic effects. In addition, number of branches per
plant, grain yield, harvest index, ascochyta blight, chocolate spot disease score and rust
disease score had high genetic advances as percent of mean. These results indicate that
whenever the best 5% accessions is selected as parent for a given trait, genotypic value of the

41
new population for the traits will be improved highly. Solieman and Ragheb (2014) had also
reported high genetic advance as percent of mean for pods per plant, total yield and branches
per plant. On the contrary, Abdellatif et al. (2020) reported low estimate of heritability and
genetic advance for rust disease reaction.

Medium heritability coupled with high genetic advance as percent of mean was observed for
chocolate spot disease score. Medium GAM values were obtained for traits like pod length
and days to flowering. On the contrary, the lowest genetic advance as percent of mean was
recorded for plant height. The result indicate selection of accessions based on this trait as
parent might results in low response in new population. In other ways this would indicate that
expression of this trait is governed by non-additive gene effects.

4.4. Correlation among Traits

The inter relationship among the 12 traits was estimated through correlation coefficient at
genotypic and phenotypic levels are presented in Table 4.5.

4.4.1. Phenotypic correlation of grain yield with other traits

The result of correlation analysis revealed that grain yield had highly significant (p<0.01) and
positive phenotypic correlations (rp) with plant height, pod length, number of pods per plant,
number of branches per plant, biomass yield, 100 seed weight and harvest index. These results
indicated that accessions with high plant height, pod length, number of pods per plant, number
of branches per plant, biomass yield, 100 seed weight and harvest index produce high grain
yield and vice versa. Therefore, these traits emerged as most important associates of grain
yield in faba bean. Similarly, Lal (2019) reported that grain yield had highly significant and
positive phenotypic correlations (rp) with plant height, number of pods per plant, number of
branches per plant, biomass yield, 100 seed weight and harvest index. Kumar et al. (2013)
reported highly significant and positive correlation of seed yield with number of branches per
plant, number of pods per plant, biological yield and harvest index. Verma et al. (2015) also

42
found highly significant and positive correlation of seed yield with harvest index and
biological yield. Singh et al. (2015) reported similar results for plant height.

Grain yield had highly significant (p<0.01) and negative phenotypic correlations (r p) with
ascochyta blight, chocolate spot and rust disease score. These results indicate that as disease
severity increase grain yield becomes highly reduced. In addition, grain yield had significant
(p<0.05) and negative phenotypic correlations (rp) with days to flowering. This result showed
that the early flowering accessions have high yield potential than those of late flowering ones.
Nchimbi and Mduruma, (2007) and Onder (2013) also reported negative association of days
to flowering with grain yield in common bean.

4.4.2. Phenotypic correlation among other traits

Rust disease score showed highly significant (p<0.01) and positive correlations with
ascochyta blight and chocolate spot, indicating that diseases are complementary to each other.
Rust disease had highly significant negative correlations with plant height, pod length,
number of pods per plant, number of branches per plant, biomass yield and100 seed weight.
This indicates that rust is negatively associated with these traits. Chocolate spot had highly
significant and positive correlations with ascochyta blight and highly significant negative
correlations with plant height, pod length, number of pods per plant, biomass yield and100
seed weight. Ascochyta blight had highly significant and negative correlations with plant
height, pod length, biomass yield and100 seed weight and significant negative correlations
with number of pods per plant. Generally the results suggested that disease related traits were
complementary to each other and negatively affected growth and yield related traits.

Harvest index showed highly significant (p<0.01) positive phenotypic correlations (r p) with
number of pods per plant. This showed that accessions with high number of pods per plant
producing high harvest index. However, harvest index had highly significant negative
phenotypic correlations with biomass yield and then significant negative phenotypic
correlations with days to flowering. Negative correlations arise due to competition among
traits for common precursors which are having restricted supply (Madhur and Jinks, 1994).

43
Hundred seed weight had high significant (p<0.01) positive phenotypic correlations with plant
height, pod length, biomass yield and significant (p<0.05) positive phenotypic correlations
with number of pods per plant. Days to flowering had high significant negative phenotypic
correlations with 100 seed weight. Similarly, Alghamdi (2007) and Tafere Mulualem et al.
(2013) reported a positive and significant phenotypic correlation between pod length and
thousand seed weight. Similar results were also obtained by Lal (2019) that hundred seed
weight had high significant positive phenotypic correlations with pod length and biomass
yield.

Biomass yield had highly significant (p<0.01) positive phenotypic correlations (r p) with plant
height, pod length, number of pods per plant and number of branches per plant. These traits
had positive correlation with biomass yield which augurs well for providing correlated
response during selection for improving biomass yield. These observations of positive
associations between biological yield and plant height, pod length, number of pods per plant
and number of branches per plant are in agreement with the reports made by Kumar et al.
(2013), Singh et al. (2015), Verma et al. (2015) and Kumar et al. (2017) on faba bean. Similar
results were obtained by Lal (2019) that biomass yield had highly significant positive
phenotypic correlations with plant height, number of pods per plant and number of branches
per plant.

Positive and highly significant (p<0.01) phenotypic correlation was recorded between number
of branches per plant with days to flowering, plant height, pod length, number of pods per
plant. Similarly, Lal (2019) obtained number of branches per plant had highly significant and
positive phenotypic correlations with plant height and number of pods per plant.

Number of pods per plant showed highly significant and positive correlation with plant height
and pod length. Thus, correlation helps breeders to identify the traits that could be used as
selection criteria in breeding program. These results suggested that improvement of grain
yield in faba bean is linked with these traits (number of pods per plant, pod length and plant
height) and selection of these traits might have good impact on yield. Azarpour et al. (2012),
Sharifi (2014) and Abdalla et al. (2015) reported positive and significant phenotypic

44
correlation of number of pods per plant with plant height. Pod length shows highly significant
(p<0.01) positive phenotypic correlations with plant height. Plant height negatively correlated
with days to flowering.

4.4.3. Genotypic correlation of grain yield with other traits

In the present study, grain yield had highly significant (p<0.01) and positive genotypic
correlations (rg) with plant height, pod length, number of pods per plant, number of branches
per plant, biomass yield, 100 seed weight and harvest index. These results indicated that
accessions with high plant height, pod length, number of pods per plant, number of branches
per plant, biomass yield, 100 seed weight and harvest index produce high grain yield. Singh et
al. (2017) reported that seed yield had highly significant and positive genotypic correlations
with number of pods per plant, 100 seed weight and harvest index. Similar result was reported
by Zakira et al. (2010) for harvest index in field pea. Grain yield had highly significant
(p<0.01) and negative genotypic correlations with ascochyta blight, chocolate spot and rust
disease score. This indicates disease plays a vital role in reduction of grain yield.

4.4.4. Genotypic correlation among other traits

Days to flowering was positively and highly significant (p<0.01) with number of branches per
plant. The result shows late flowering accessions are expected to produce high number of
branches per plant. Days to flowering negatively correlated with100 seed weight and harvest
index. This result showed that the late flowering accessions have less yield potential than
those of early flowering ones. These findings are in concurrence with Alghamdi (2007), Tofiq
et al. (2016). Kumar et al. (2020) also reported days to flowering negatively correlated with
100 seed weight.

Plant height had positive and high significant (p<0.01) genotypic correlation with pod length,
number pods per plant, number of branches per plant, biomass yield and 100 seed weight.
These indicate accessions that have high plant height had high pod length, number of pods per
plant, number of branches per plant, biomass yield and 100 seed weight. Similar results

45
obtained by Lal (2019) number of branches per plant, biomass yield positively and high
significant correlation with plant height on faba bean. Plant height had negative and high
significant (p<0.01) correlation with chocolate spot and rust disease score and negatively
correlated with ascochyta blight. This indicates disease plays a vital role in reduction of plant
height.

Pod length had positive and high significant (p<0.01) correlation with number of branches per
plant, biomass yield and 100 seed weight and positive significant correlation with number of
pods per plant. Lal, (2019) reported that pod length had positive and high significant
correlation with 100 seed weight. Pod length had negative and high significant correlation
with ascochyta blight, chocolate spot and rust disease score. Number of pods per plant had
positive and high significant (p<0.01) correlation with number of branches per plant, biomass
yield and positive and significant correlation with harvest index. Improving these traits
increases the development of number of pods per plant that support to increases grain yield.
This result agrees with Singh et al. (2017) also reported number of pods per plant had positive
and high significant correlation with number of branches per plant and harvest index. Similar
result also reported by (Million Fikreselassie, 2012). Number of pods per plant had negative
and high significant (p<0.01) correlation with chocolate spot and rust disease score. Number
of branches per plant had positive and significant correlation with biomass yield and negative
highly significant correlation with rust disease score.

Biological yield had positive and high significant (p<0.01) genotypic correlation with 100
seed weight. This indicates that breeding for increased biological yield might result in high
100 seed weight. This implies that the trait has positive impact on faba bean improvement.
Biological yield had negative high significant (p<0.01) genotypic correlation with harvest
index, chocolate spot and rust disease score and negative significant correlation with
ascochyta blight. This implies that these traits inversely correlated with biological yield.
Hundred seed weight had negative high significant correlation with ascochyta blight and
chocolate spot but negative significant correlation with rust. These indicate that hundred seed
weight negatively affected by disease and leads to low yield. Ascochyta blight had positive
high significant correlation with chocolate spot and rust diseases score and also chocolate spot

46
had positive high significant correlation with rust disease. This result indicates the occurrence
of one disease simply favored another.

47
 Table 4.5. Estimates of genotypic (below diagonal) and phenotypic (above diagonal) correlation coefficients of 12 traits of 81 faba
bean accessions

Traits DF PH PL NPP NBP BY GY HSW HI AB CS RT

DF 1 -0.198* -0.066ns -0.089ns 0.293** 0.016ns -0.179* -0.248** -0.192* 0.072ns 0.132ns 0.102ns
PH -0.178ns 1 0.542** 0.588** 0.333** 0.456** 0.576** 0.491** 0.111ns -0.206** -0.412** -0.398**
PL -0.071ns 0.560** 1 0.287** 0.280** 0.390** 0.360** 0.709** -0.026ns -0.239** -0.289** -0.263**
NPP -0.078ns 0.628** 0.271* 1 0.446** 0.518** 0.755** 0.175* 0.228** -0.186* -0.348** -0.394**
NBP 0.381** 0.369** 0.312** 0.454** 1 0.229** 0.336** 0.109ns 0.077ns -0.028ns -0.092ns -0.248**
BY 0.026ns 0.524** 0.438** 0.538** 0.238* 1 0.562** 0.396** -0.458** -0.244** -0.505** -0.561**
ns
GY -0.190 0.680** 0.407** 0.789** 0.374** 0.579** 1 0.352** 0.418** -0.281** -0.483** -0.413**
HSW -0.252* 0.510** 0.789** 0.136ns 0.061ns 0.418** 0.362** 1 -0.009ns -0.301** -0.411** -0.263**
HI -0.227* 0.173ns -0.007ns 0.272* 0.108ns -0.427** 0.438** -0.009ns 1 0.003ns -0.032ns 0.189*
AB 0.075ns -0.247* -0.289** -0.206ns -0.041ns -0.255* -0.286** -0.329** 0.004ns 1 0.339** 0.332**
CS 0.142ns -0.516** -0.373** -0.373** -0.116ns -0.514** -0.499** -0.458** -0.060ns 0.348** 1 0.413**
RT 0.105ns -0.495** -0.332** -0.429** -0.294** -0.578** -0.422** -0.283** 0.199ns 0.338** 0.428** 1

Note: ** = Highly significant (p<0.01), * = Significant (p<0.05), ns = Non significant, DF= days to flowering, PH= plant height, PL= pod
length, NPP= number of pods per plant, NBP= number of branch per plant, BY= biomass yield, GY= grain yield, HSW= 100 seed weight, HI=
harvest index, AB= ascochyta blight, CS= chocolate spot, RT= rust disease score.

48
4.5. Direct and Indirect Effects of Traits

The phenotypic and genotypic direct and indirect effects of different traits on grain yield are
presented in Table 4.6. and 4.7., respectively.

4.5.1. Phenotypic path coefficients

In the present study, the path coefficient analysis was carried out by using phenotypic
correlation coefficients among twelve traits. The high positive direct effects on grain yield
were exerted by biological yield (0.856) and harvest index (0.778) (Table 4.6.). The results
indicate biological yield and harvest-index are most important traits to which emphasis should
be given during simultaneous selection aimed at improving grain yield in faba bean. These
results are in close association with previous workers (Salem, 1982; Bora et al., 1998; Kumar
et al., 2013) for biological yield and harvest-index traits on faba bean. Similar result was also
obtained by Lal, (2019) on faba bean for traits biological yield and harvest-index.

Biological yield exerted considerable positive indirect effects on grain yield via number of
pods per plant, plant height, pod length and 100 seed weight. Harvest index exhibited high
positive indirect effects on grain yield via number of pods per plant and plant height. These
results suggested the importance of these traits in selection for the improvement of grain
yield. However, biological yield exerted considerable negative indirect effects on grain yield
via harvest index chocolate spot and rust disease score and harvest index exhibited high order
of negative indirect effects on grain yield via biological yield. Similarly, Lal (2019) observed
biological yield exerted considerable positive indirect effects on grain yield via number of
pods per plant, number of branches per plant and plant height. Harvest index exhibited high
order of positive indirect effects on grain yield via number of pods per plant and biological
yield on faba bean. In addition to the most important direct yield contributors, biological yield
and harvest index had considerable positive indirect effects via different traits are also most
important indirect yield contributors, implying that biological yield and harvest index are
important traits to be considered during devising selection strategy aimed at developing high

49
yielding varieties in faba bean. Plant height, pod length, number of pods per plant and number
of branches per plant had positive direct effects on grain yield. Ulukan et al. (2003) and
Tadele Tadesse et al. (2011) also observed the highest positive direct effect of number of pods
per plant together with plant height. Kumar et al. (2017) also reported that the number of
branches per plant, number of pods per plant and pod length exhibited positive direct effect on
yield. On the contrary, days to flowering, 100 seed weight, ascochyta blight and rust disease
score had negative direct effects on grain yield. Tofiq et al. (2016) also reported negative
direct effect of 100 seeds weight on yield.

The magnitude of residual effect (0.297) indicated that the traits included in the study
accounted for most of the variability present in grain yield, indicating that the contribution of
traits considered was 70.3% and the rest 29.7% was the contribution of other traits which
were not considered in the present study.

4.5.2. Genotypic path coefficients

The high positive direct effects on grain yield were exerted by biological yield (0.844) and
harvest index (0.756) (Table 4.7.). Thus, biological yield and harvest-index emerged as most
important direct yield components on which emphasis should be given during simultaneous
selection aimed at improving grain yield in faba bean. These results are in close association
with those of Salem, (1982), Bora et al. (1998) and Kumar et al. (2013). Similar results also
obtained by Lal (2019). The direct effects of remaining traits were low to be considered
important. Biological yield exerted considerable high positive indirect effects on grain yield
via number of pods per plant, plant height, pod length and 100 seed weight but biological
yield exerted considerable high negative indirect effects on grain yield via harvest index,
chocolate spot and rust disease score. Harvest index exhibited high order of positive indirect
effects on grain yield via number of pods per plant and plant height but harvest index
exhibited high order of negative indirect effects on grain yield via biological yield. Similar
results were obtained by Lal, (2019) for biological yield and harvest index. In addition to
emerging as most important direct yield contributors owing to their very high positive direct
effects on grain yield, biological yield and harvest index, having considerable positive indirect

50
effects via different traits, also appeared as most important indirect yield components. The
indirect effects of remaining traits were too low to be considered important.

In the present study, path analysis identified biological yield followed by harvest-index as
most important direct as well as indirect yield contributing traits. These indicate that both
traits had true association with grain yield and their importance in determining these complex
traits. Therefore, important consideration should be given while practicing selection aimed at
the improvement of grain yield in faba bean. These results were in accordance with the result
of Jivani et al. (2013) who reported the highest direct effect of harvest index and biological
yield on grain yield in chickpea. Plant height, number of pods per plant and number of
branches per plant had positive direct effects on grain yield at genotypic level but days to
flowering, pod length, 100 seed weight, ascochyta blight and rust disease score had negative
direct effects on grain yield.

The residual effect shows how much the explanatory variables represent the variability of the
dependent variable (Singh and Chaudhary, 1985). The residual effect in present study at the
genotypic path coefficient was 0.272; as a result, the studied traits explain 72.8 % of the
variability in the seed yield and show that we did not consider few traits which are related to
grain yield.

51
Table 4.6. Phenotypic direct (bold and diagonal) and indirect effects of 11 traits on grain yield of 81 faba bean accessions

Traits DF PH PL NPP NBP BY HSW HI AB CS RT

DF -0.052 -0.007 -0.001 -0.008 0.013 0.014 0.011 -0.149 -0.005 0.006 -0.002
PH 0.010 0.034 0.006 0.053 0.014 0.390 -0.022 0.087 0.013 -0.019 0.008
PL 0.003 0.019 0.011 0.026 0.012 0.333 -0.032 -0.020 0.016 -0.013 0.005
NPP 0.005 0.020 0.003 0.090 0.019 0.443 -0.008 0.178 0.012 -0.016 0.008
NBP -0.015 0.011 0.003 0.040 0.043 0.196 -0.005 0.060 0.002 -0.004 0.005
BY -0.001 0.016 0.004 0.047 0.010 0.856 -0.018 -0.356 0.016 -0.023 0.012
HSW 0.013 0.017 0.008 0.016 0.005 0.339 -0.045 -0.007 0.020 -0.019 0.005
HI 0.010 0.004 0.000 0.021 0.003 -0.392 0.000 0.778 0.000 -0.001 -0.004
AB -0.004 -0.007 -0.003 -0.017 -0.001 -0.209 0.014 0.003 -0.065 0.015 -0.007
CS -0.007 -0.014 -0.003 -0.031 -0.004 -0.432 0.018 -0.025 -0.022 0.045 -0.009
RT -0.005 -0.014 -0.003 -0.035 -0.011 -0.480 0.012 0.147 -0.022 0.019 -0.021

Residual factor = 0.297

Note: DF= days to flowering, PH= plant height, PL= pod length, NPP= number of pods per plant, NBP= number of branch per plant, BY=
biomass yield, HSW= 100 seed weight, HI= harvest index, AB= ascochyta blight, CS= chocolate spot, and RT= rust disease score.

52
Table 4.7. Genotypic direct (bold and diagonal) and indirect effects of 11 traits on grain yield for 81 faba bean accessions

Traits DF PH PL NPP NBP BY HSW HI AB CS RT

DF -0.061 -0.011 0.001 -0.005 0.026 0.022 0.008 -0.172 -0.005 0.009 -0.002
PH 0.011 0.061 -0.006 0.042 0.025 0.442 -0.017 0.131 0.017 -0.034 0.008
PL 0.004 0.034 -0.010 0.018 0.021 0.370 -0.026 -0.005 0.020 -0.024 0.006
NPP 0.005 0.038 -0.003 0.066 0.031 0.454 -0.004 0.206 0.014 -0.024 0.007
NBP -0.023 0.022 -0.003 0.030 0.067 0.201 -0.002 0.082 0.003 -0.008 0.005
BY -0.002 0.032 -0.004 0.036 0.016 0.844 -0.014 -0.323 0.017 -0.034 0.010
HSW 0.015 0.031 -0.008 0.009 0.004 0.353 -0.032 -0.007 0.023 -0.030 0.005
HI 0.014 0.010 0.000 0.018 0.007 -0.361 0.000 0.756 0.000 -0.004 -0.003
AB -0.005 -0.015 0.003 -0.014 -0.003 -0.215 0.011 0.003 -0.068 0.023 -0.006
CS -0.009 -0.031 0.004 -0.025 -0.008 -0.434 0.015 -0.046 -0.024 0.066 -0.007
RT -0.006 -0.030 0.003 -0.029 -0.020 -0.488 0.009 0.150 -0.023 0.028 -0.017

Residual factor = 0.272

Note: DF= days to flowering, PH= plant height, PL= pod length, NPP= number of pods per plant, NBP= number of branch per plant, BY=
biomass yield, HSW= 100 seed weight, HI= harvest index, AB= ascochyta blight, CS= chocolate spot, RT= rust disease.

53
 4.6. Clustering among Accessions

Cluster analysis distinguished the 81 accessions into seven groups of di fferent sizes (Table
4.8. and Figure 4.1.), members within a cluster being assumed to be more closely related in
terms of the traits under consideration with each other than those members in di fferent
clusters. Similarly, members in clusters with non-significant distance were assumed to have
more close relationships with each other than they are with those in significantly distant
clusters. The number of accessions in each of the seven clusters ranged from 3 to 24 in the
smallest and largest cluster, respectively.

Table 4.8. List of faba bean accessions grouped into seven clusters

Cluste
r Number of Name of accessions

accessions

CI 8 212565, 212568, 25018, 212567, 208085, 212575, 25010, 215748

CII 9 245140, 212566, 229303, 219355, 212811, 25003, 25323, 213214, 228607

CIII 3 Tumsa, Dosha, Local

CIV 24 27279, 25006, 27052, 203105, 25279, 208114, 25022, 25280,235956,

220079, 25277, 220076, 25264,212580, 212578, 25338, 226125,

25346, 25340, 215128, 25341, 25329, 25332, 25333

CV 12 25274, 229310, 25270, 25290, 235709, 219089, 240497, 25337,

235433, 25325, 25334, 25310
CVI 8 212572, 212576, 27290, 25336, 25328, 25339, 215129, 25331

CVII 17 25299, 25298, 235955, 25292, 25017, 213211, 25335, 25309, 25304,
25330, 25307,25306, 25311, 25302, 25327, 25303, 25301

Clusters CI and CVI had 8 accessions for each. Clusters CII and CIII had 9 and 3 accessions,
respectively. Cluster CIV was the largest cluster group consisting of 24 accessions followed

54
by CVII which had 17 accessions. CV had 12 accessions. These indicate the presence of
adequate variability among the accessions. The discrimination of accessions into so many
discrete clusters suggested presence of genetic diversity in the material evaluated. Presence of
substantial genetic diversity among the accessions implied that these accessions may serve as
good source for selecting the diverse parents for hybridization program aimed at isolating
desirable segregants for grain yield and other important traits. Similarly, substantial genetic
divergence in the faba bean germplasm was reported previously (Nanda et al., 1991;
Gemechu Keneni et al., 2005; Qi and Xing, 2009; Wang et al., 2012; Berma et al., 2016 and
Chaudhary et al., 2018).

55
Figure 4.2. Dendrogram showing relationships among 81 Ethiopian faba bean accessions

56
4.6.1. Genetic distance among clusters

Pair wise generalized distances (D2) among the seven clusters were presented in Table 4.9.
Out of 21 possible pairs of clusters, differences between 14 pairs were highly significant
(P<0.01) and 4 pairs were significant (P<0.05), while those between the rest of the clusters
were non-significant. The maximum inter cluster distance (D 2 = 152.28**) was detected
between CI and CIII. Cluster CI constituted accessions from two regions of semen Shewa
(Amhara and Oromia) except one while cluster CIII constituted standard and local checks as
clearly shown from appendix Table 1 of the 81 Faba bean accessions used in the study. The
high values of inter cluster distances indicated divergence among the accessions and might be
used in breeding programs for better genetic recombination and selection of genetically
divergent parents for exploitation in crossing programs. This finding is consistent with
Million Fikreselassie and Habtamu Seboka (2012) and Gemechu Keneni et al. (2005) who
used 25 and 160 faba bean genotypes respectively and found high D 2 value. Gemechu Keneni
et al. (2007) also reported divergence between Ethiopian faba bean germplasms. The second
most divergent clusters were cluster CIII and CIV (D 2 = 130.22**). Cluster CIV constituted
accessions from all regions (Amhara, Oromia, Tigray and Southern nation nationalities
people) (appendix Table 1). This indicates accessions from different regions might have
similar genetic background.

The third most divergent clusters were CIII and CVI (D 2 = 121.99**). Cluster CVI constituted
accessions from three regions (Amhara, Oromia and Southern nation nationalities people).
The results implied that crossing of accessions from CIII with accessions from CI, CIV and
CVI will result in useful segregants in the F1 and subsequent generations. Similar findings
were reported by Lal et al. (2019) who clustered 83 faba bean genotypes in to ten clusters.
The minimum distance (D2 = 6.93) was observed between CIV and CVII suggesting that the
materials may be very closely related. Cluster CVII constituted accessions from Southern
nation nationalities people region except three accessions. In the present investigation, the
highest intra-cluster distance was found for CI followed by CIV, CII, CV, CVII, CVI and
CIII. This implies that the accessions present in a cluster have little genetic divergence from

57
each other with respect to aggregate effect of twelve traits under study, while more genetic
diversity was observed between the accessions belonging to different clusters.

Table 4.9. Estimates of average intra (diagonal) and inter (above diagonal) cluster distances
for 7 clusters of 81 faba bean accessions

Cluste
CI CII CIII CIV CV CVI CVII
r

206.9 27.25** 152.28** 19.54 58.57** 49.81** 29.65**

CI
9
CII 170.43 104.81** 13.92 22.94* 37.03** 19.69*
CIII 10.90 130.22** 109.29** 121.99** 93.40**
CIV 195.76 23.29* 24.57* 6.93
CV 91.32 29.22** 25.57**
CVI 27.21 26.11**
CVII 61.58
** = highly significant at probability level p<0.01 (χ 2 = 24.72), * = Significant at probability level of
p<0.05 (χ2=19.67)

4.6.2. Mean performance of clusters

Cluster means showed significant differences among the clusters for most of the traits (Table
4.10.). Cluster CI constituted inferior accessions (susceptible to chocolate spot, rust, the
shortest in plant height, low 100 seed weight and harvest index) for most of the traits. Similar
results were obtained by Gemechu Keneni et al. (2005) for chocolate spot, the shortest in
plant height, and low 100 seed weight. Clusters CII, CIV and CV comprised intermediate
accessions for most of the traits, but CV superior in grain yield. Cluster CVI showed the
highest mean for harvest index and lowest mean of biological yield, ascochyta blight score
and rust disease score. Cluster CVII was characterized by the least days to flowering, number
of pods per plant and number of branches/plant but superior in ascochyta blight disease.
Cluster CIII constituted superior accessions for most of the traits including chocolate spot

58
disease, rust disease, plant height, pod length, number of pods per plant, number of branches
per plant, biological yield and 100 seed weight.

Overall, the results showed wide variation from one cluster to another in respect of cluster
means for twelve traits, which indicated that accessions having distinctly different mean
performance for various traits were separated into different clusters. The crossing between the
entries belonging to cluster pairs having large intercluster distance and possessing high cluster
means for one or other traits to be improved may be recommended for isolating desirable
recombinants in the segregating generations in faba bean. However, the results suggested that
selection of parents should consider not only the distance between clusters but also the special
merits of each cluster and each accession within a cluster depending on the specific objectives
of hybridization as suggested by Singh (1990).

Table 4.10. Cluster means for 12 traits of 81 faba bean accessions

Traits CI CII CIII CIV CV CVI CVII Mean

DF 50.88a 49.72 45.17 44.71 44.58 43.94 43.56b 46.08

PH 84.04b 103.52 121.71a 93.01 109.02 91.42 98.63 100.19
PL 3.46 3.84 5.78a 3.44b 3.75 3.54 4.04 3.98
NPP 9.47 10.30 19.37a 9.75 16.53 13.03 8.86b 12.47
NBP 1.78 2.18 2.40a 1.55 2.06 1.45 1.35b 1.82
9926.0 8145.6
BY 7283.69 4 13349.67a 7520.44 10402.26 5211.28b 3 8834.14
1979.3 1910.4
GY 1438.02b 6 2962.10 1703.18 3231.91a 1958.92 7 2169.14
HSW 33.29b 38.13 82.32a 37.92 42.38 40.06 46.76 45.83
HI 20.22b 20.65 22.52 23.58 32.10 38.98a 23.80 25.98
AB 4.25 3.78 2.33 3.38 2.33 4.50a 2.29b 3.27
CS 5.25a 2.11 1.00b 3.75 1.67 3.00 2.82 2.80
RT 4.63a 1.00b 1.00b 2.17 1.17 4.63a 2.35 2.42
Note: a=Highest value, b=Lowest value.

59
DF= days to flowering, PH= plant height, PL= pod length, NPP= number of pods per plant, NBP=
number of branch per plant, BY= biomass yield, GY= grain yield, HSW= 100 seed weight, HI=
harvest index, AB= ascochyta blight, CS= chocolate spot, RT= rust disease score.

4.7. Principal Component Analysis

In the present study, the first four principal components exhibited eigen values more than one
(4.856, 2.006, 1.672 and 1.063) and explained about 73.81% of the total variations among the
81 faba bean accessions (Table 4.11.). The first four PCs were given due importance for
further explanation. Similar findings were reported by Tiwari (2019) who found the first three
principal components exhibiting eigen values more than one and explained about 75.53% of
the total variations.

The first principal component accounted for 37.35% of the total variations. Traits such as
plant height, pod length, number of pods per plant, biological yield, grain yield and 100 seed
weight had high contribution to the first PC. These indicated that these traits had higher
relative contribution to the total diversity and they were the ones that most di fferentiated the
accessions. Gemechu Keneni et al. (2005) reported traits with relatively greater weight in PC1
like grain yield and plant height had higher relative contribution to the total diversity and they
were the ones that most differentiated the populations. Similarly, the high contribution of
number of pods per plant and grain yield in the first principal component was reported by
Kumar et al. (2019). The second component accounted for 15.43% of the total variation and
predominantly illustrates variation in days to flowering and number of branches per plant. The
third principal component accounted for 12.86% of the total variation and it was chiefly
accounted by variation in number of pods per plant, grain yield and harvest index. The fourth
principal component accounted for 8.17% of the total variation and indicated with high
variation in pod length, number of branches per plant, 100 seed weight and rust disease score.

60
Table 4.11. Eigenvectors, eigen values and variance explained by the first four principal
components

Traits PC-1 PC-2 PC-3 PC-4

DF -0.090 0.537 -0.152 0.285

PH 0.363 0.113 0.102 0.061
PL 0.325 -0.099 -0.156 0.542
NPP 0.322 0.221 0.307 -0.264
NBP 0.184 0.481 0.162 0.302
BY 0.333 0.171 -0.324 -0.247
GY 0.373 0.055 0.325 -0.123
HSW 0.303 -0.285 -0.213 0.463
HI 0.046 -0.139 0.706 0.180
AB -0.208 0.087 0.138 0.071
CS -0.313 0.050 0.081 0.098
RT -0.304 -0.094 0.202 0.344
Eigen value 4.856 2.006 1.672 1.063
Proportion (%) 37.35 15.43 12.86 8.17
Cumulative (%) 37.35 52.78 65.64 73.82

Note: DF= days to flowering, PH= plant height, PL= pod length, NPP= number of pods per plant,
NBP= number of branch per plant, BY= biomass yield, GY= grain yield, HSW= 100 seed weight, HI=
harvest index, AB= ascochyta blight, CS= chocolate spot, RT= rust disease.

As it is visualized in Figure 4.2., standard checks and local genotype in the present study were
found in closed distance in the quadrants of the principal component axis. Treatment 8 and 16
was plotted far apart from others. Treatment number 8, 16, 7, 45, 15 and 29 highly contributed
to PC1 in negative direction and Treatment 62, 63, 78, and 81 contributed highly in positive
direction for PC1. The biplot gave more opportunity to assess which accessions were good for
which traits that would help as excellent baseline information for faba bean improvement.
Based on this treatment 4, 10, 14, 21, 23, 24, 25, 36, 41, 42, 43, 52, 58, 62, 63, and 64 can
directly improve by select grain yield whereas, biomass yield well for 21 and 58. As a result,

61
grain yield was highly correlated with number of pods per plant, biomass yield and plant
height, whereas weaker but positive correlated with number of branches per plant while
negatively correlated with ascochyta blight, chocolate spot and rust disease score. Treatment
28, 29, 31, 33, 35, 3, 19, 18, 45, 54, 56, 48, 57, 47, 50, 75, 13, 44, 53, 55, 61, 76 negatively
correlated with grain yield and also had low yields. This implies a substantial amount of
diversity in the faba bean accessions and possibility of its improvement.

Figure 4.3. Principal component analysis biplot describing the relative position of 81 faba
bean accessions and 12 traits.

62
 Chapter 5. CONCLUSION AND RECOMMENDATIONS

5.1. Conclusion

The analysis of variance showed highly significant differences (p<0.01) among accessions for
all traits except days to maturity and number of seeds per pod. This indicates the existence of
variability among accessions for most studied traits. High (>20%) genotypic and phenotypic
coefficients of variation were observed for 100 seed weight, biomass yield, harvest index,
grain yield, number of branches per plant, number of pods per plant, ascochyta blight,
chocolate spot and rust disease score. These indicate the existence of wide genetic variation
among the accessions and showed the possibility of genetic improvement of these traits
through selection

In the present study, high estimates of heritability (>60%) were found for days to flowering,
number of pods per plant, biomass yield, grain yield, 100 seed weight, harvest index and
ascochyta blight, chocolate spot and rust diseases score. The high H 2 values of these traits
indicate possibility of improving these traits through selection. High heritability coupled with
high genetic advance as percent of the mean were observed for hundred seed weight, harvest
index, number of pod per plant, grain yield, biological yield, ascochyta blight, chocolate spot
and rust disease. These results indicated that those traits are governed by additive gene action.
Therefore, these traits could be easily improved through direct selection.

Grain yield had highly significant (p<0.01) and positive phenotypic and genotypic correlation
with plant height, pod length, number of pods per plant, number of branches per plant,
biological yield, 100-seed weight and harvest index. These results indicated the possibility of
simultaneous improvement of grain yield with these traits through selection. Path coefficient
analysis showed that harvest index and biological yield had the highest direct effects on yield
both at phenotypic and genotypic levels, indicating the importance of these traits for indirect
selection of faba bean accessions for simultaneous improvement of grain yield through
biological yield and harvest index.

63
Cluster analysis grouped the accessions into seven clusters based on their similarity and
dissimilarity. The maximum inter cluster distance (D 2 = 152.28**) was detected between CI
and CIII followed by CIII-CIV and CIII-CVI. The high values of inter cluster distances
indicated divergence among the accessions and might be used in breeding programs for better
genetic recombination and selection of genetically divergent parents for exploitation in
crossing programs. The principal component analysis show the first four principal
components exhibited eigen values more than one and explained about 73.81% of the total
variations. Generally, it can be concluded that there was considerable variations among faba
bean accessions for the traits considered, implying the possibility of improving these traits
through direct and indirect selection.

5.2. Recommendations

Based on the results of the present study, the following recommendations are made for future
consideration:
 The results revealed the presence of an immense amount of genetic variation among
Ethiopian faba bean accessions that breeders could consider future faba bean breeding
program.
 However, these genetic resources should be further characterized over seasons and
locations and using molecular tools.

64
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7. APPENDICES

85
 Appendix Table 1. List of the 81 faba bean accessions used in the study

TRT Accession No Region/State Zone Woreda/District Altitude Coll.Date Local Name

1 212565 Amhara Semen Shewa Siyadebrina Wayu Ens 2670 30-Nov-84 Bakela

2 27279 Amhara Misrak Gojam Machakel 2400 13-Dec-81 Bakela

3 25299 SNNP Kembata Alabana Temb Angacha 2220 05-Dec-78 Bakela
4 245140 SNNP Gedeo Dila Zuria 2489 02-Jan-08 Bakela
5 212566 Oromiya Semen Shewa Wuchalena Jido 2630 30-Nov-84 Bakela
6 25006 Amhara Misrak Gojam Hulet Ej Enese 2450 26-Nov-77 Bakela
7 212568 Amhara Semen Shewa Siyadebrina Wayu Ens 2670 01-Dec-84 Bakela
8 25018 Oromiya Semen Shewa Wuchalena Jido 2440 04-Dec-77 Bakela
9 27052 Oromiya Arssi Kofele 2300 14-Jan-79 Bakela
10 25274 Oromiya Mirab Shewa Becho 1830 01-Dec-78 Bakela
11 229303 Amhara Semen Shewa Lay Betna Tach Bet 2340 18-Nov-88 Bakela

12 212567 Amhara Semen Shewa Siyadebrina Wayu Ens 2630 30-Nov-84 Bakela
13 25298 SNNP Kembata Alabana Temb Angacha 2420 05-Dec-78 Bakela
14 219355 Oromiya Borena Adolana Wadera 1730 26-Jul-86 Ater

15 208085 Amhara Semen Shewa Weremo Wajetuna Mida 2300 25-Nov-83 Bakila

16 212811 Amhara Semen Gondar Wegera 2900 20-Dec-84 Bakela

17 203105 Oromiya Illubabor Dedesa 2170 12-Dec-82 Bakela

18 235955 Amhara Semen Gondar Debark 2940 15-Dec-93 Bakela

19 25279 SNNP Gurage Cheha 2980 02-Dec-78 Bakela

20 208114 Amhara Semen Shewa Weremo Wajetuna Mida 1983 29-Nov-83 Bakila

21 229310 Amhara Semen Shewa Weremo Wajetuna Mid 2140 21-Nov-88 Bakela

86
TRT Accession No Region/State Zone Woreda/District Altitude Coll.Date Local Name

22 212572 Amhara Semen Shewa Weremo Wajetuna Mid 2630 02-Dec-84 Bakela
23 25003 Oromiya Semen Shewa Kuyu 2490 03-Dec-77 Bakela
24 25323 SNNP Gurage Selti 2110 07-Dec-78 Bakela
25 25022 Oromiya Semen Shewa Kuyu 2480 22-Nov-77 Bakela
26 25280 SNNP Gurage Cheha 3070 02-Dec-78 Bakela

27 235956 Amhara Semen Gondar Debark 2870 15-Dec-93 Bakela

28 220079 Tigray Mehakelegnaw Adwa 2750 21-Nov-86 Ate bahiri

29 212576 Amhara Semen Shewa Lay Betna Tach Bet 2610 03-Dec-84 Bakela
30 212575 Amhara Semen Shewa Lay Betna Tach Bet 2600 03-Dec-84 Bakela
31 25277 SNNP Gurage Cheha 2870 02-Dec-78 Bakela

32 220076 Tigray Mehakelegnaw Adwa 1920 17-Nov-86 Atebahri

33 27290 Oromiya Misrak Wellega Jimma Arjo 2280 18-Jan-82 Bakelem

34 25264 Oromiya Mirab Shewa Becho 2350 29-Nov-78 Bakela
35 25292 SNNP Hadiya Limo 2325 04-Dec-78 Bakela
36 25270 Oromiya Mirab Shewa Becho 2000 29-Nov-78 Bakela

37 212580 Amhara Semen Shewa Mama Midrina Lalo Mi 3130 05-Dec-84 .

38 25017 Amhara Misrak Gojam Enarj Enawga 2370 25-Nov-77 Bakela

39 212578 Amhara Semen Shewa Geramidirna Keya Geb 3100 05-Dec-84 Bakela
40 25010 Oromiya Semen Shewa Gerar Jarso 2870 28-Nov-77 Bakela
41 25290 SNNP Hadiya Limo 2790 03-Dec-78 Bakela
42 235709 SNNP Bench Maji DIRASHE Special 2150 23-Aug-93 Bakela
43 219089 Oromiya Arssi Amigna 2410 05-Jul-86 Bakela
44 25338 SNNP Gurage Meskanena Mareko 08-Dec-78 Bakela

87
TRT Accession No Region/State Zone Woreda/District Altitude Coll.Date Local Name
45 215748 Amhara Debub Wello Dessie Zuria 07-Dec-85 .
46 213211 Oromiya Jimma Tiro Afeta 06-Apr-77 Bakella
47 25336 SNNP Gurage Meskanena Mareko 07-Dec-78 Bakela
48 226125 Amhara Debub Wello Legambo 14-Jan-88 Bakella

49 240497 SNNP Keficho Shekicho Decha 23-Jan-00 Bakello

50 25346 SNNP Gurage Meskanena Mareko 08-Dec-78 Bakela
51 25328 SNNP Gurage Selti 07-Dec-78 Bakela
52 213214 Oromiya Jimma Limu Seka 24-Apr-77 Bakella
53 25335 SNNP Gurage Meskanena Mareko 07-Dec-78 Bakela
54 25340 SNNP Gurage Meskanena Mareko 08-Dec-78 Bakela
55 25339 SNNP Gurage Meskanena Mareko 08-Dec-78 Bakela
Kemer
56 215129 Amhara Semen Shewa Mama Midrina Lalo Mi 22-Dec-84 Dengay

57 215128 Amhara Semen Shewa Mama Midrina Lalo Mi 25-Dec-84 Kimi dingay

58 228607 Amhara Misrak Gojam Goncha Siso Enese 05-May-88 Bakela

59 25337 SNNP Gurage Meskanena Mareko 08-Dec-78 Bakela
60 25341 SNNP Gurage Meskanena Mareko 08-Dec-78 Bakela
61 25331 SNNP Gurage Selti 07-Dec-78 Bakela

62 235433 Tigray Mehakelegnaw Kola Temben 05-Dec-92 Alqui

63 Tumsa
64 25325 SNNP Gurage Selti 07-Dec-78 Bakela
65 25329 SNNP Gurage Selti 07-Dec-78 Bakela
66 25309 SNNP Kembata Alabana Temb Angacha 05-Dec-78 Bakela
Bakela
67 25304 SNNP Kembata Alabana Temb Angacha 05-Dec-78

88
68 25330 SNNP Gurage Selti 07-Dec-78 Bakela

TRT Accession No Region/State Zone Woreda/District Altitude Coll.Date Local Name

69 25307 SNNP Kembata Alabana Temb Angacha 05-Dec-78 Bakela
70 25334 SNNP Gurage Selti 07-Dec-78 Bakela
71 25306 SNNP Kembata Alabana Temb Angacha 05-Dec-78 Bakela
72 25311 SNNP Kembata Alabana Temb Angacha 05-Dec-78 Bakela
73 25332 SNNP Gurage Sodo 07-Dec-78 Bakela
74 25310 SNNP Kembata Alabana Temb Angacha 05-Dec-78 Bakela
75 25333 SNNP Gurage Selti 07-Dec-78 Bakela
76 25302 SNNP Kembata Alabana Temb Angacha 05-Dec-78 Bakela
77 25327 SNNP Gurage Selti 07-Dec-78 Bakela
78 Dosha
79 25303 SNNP Kembata Alabana Temb Angacha 05-Dec-78 Bakela
80 25301 SNNP Kembata Alabana Temb Angacha 05-Dec-78 Bakela
81 Local

89
 Appendix Table 2. Mean performance of 81 faba bean accessions for 14 traits tested at Kudmi in 2019

TRT DF DM PH PL NPP NSP NBP BY GY HSW HI AB(%) CS (%) RT (%)

1 52bc 124a-f 92.9i-x 4.2c-g 8.1r-z 3a-e 2c-j 6742.4r-b 1420.1e-j 36.1n-b 21.06o-a 33.330d 55.560c 33.330d

2 46.5g-k 116.5d-m 99f-u 3.9d-l 12.4h-u 2.9a-f 1.7e-m 9353.1g-n 1669.4x-d 35.35p-b 17.85v-b 55.560b 33.330e 33.330d

3 42.5n 116.5d-m 100.3c-t 4.35c-f 7.7s-z 2.9a-f 1.6f-m 7522.6o-w 1799.3r-a 50b-g 23.935k-y 33.330d 44.445d 55.560b

4 52bc 123a-g 104.7c-o 3.7e-m 10.6l-y 3.1a-d 2c-j 11305.6c-f 1817.4q-z 37.1m-a 16.085z-b 55.560b 11.110g 11.110e

5 53b 124a-f 92.2i-y 3.95c-l 14.1f-o 2.8b-g 1.8e-l 10504.2c-i 2023.4k-r 36.25n-b 19.255s-b 44.445c 33.330e 11.110e

6 47.5f-i 113i-m 92.3i-y 2.8n 8.1r-z 2.3fg 0.6n 10195.2e-k 1285h-j 32.55s-b 12.61b 33.330d 33.330e 33.330d

7 51b-e 116e-m 71.6y 3.35g-n 11.7i-w 2.7b-g 1.5g-m 6696.4s-b 1684.8v-d 32.6s-b 25.305j-u 77.780a 66.670b 55.560b

8 59a 125a-d 75.4w-y 3.1l-n 3.95z 1h 2c-j 6666.7t-b 1305.7j-j 31.4v-b 20.855o-a 33.330d 66.670b 55.560b

9 49d-g 112.5j-m 88.6l-y 3.45g-n 9.4o-y 2.9a-f 1.6f-m 5625y-c 1478.4c-h 37.65k-a 26.715i-r 33.330d 33.330e 33.330d

10 46h-l 113i-m 97.8f-v 3.65e-n 13.8g-p 3.5a 2.2b-h 8823.5i-p 2284.7g-j 44c-r 25.895j-t 11.110f 11.110g 11.110e

11 48.5e-h 118.5c-l 83.7p-y 3.6e-n 12.8h-s 2.8b-g 1.8e-l 11477.6b-e 1798.2r-a 32.4t-b 15.665ab 33.330d 11.110g 11.110e

12 51.5bcd 119.5c-l 81s-y 3.55e-n 7.7s-z 2.6c-g 2.4a-f 8010.4m-u 1604.8y-e 31.3v-b 20.09r-a 33.330d 44.445d 55.560b

13 42n 116e-m 93.7h-x 3.9d-l 10.2m-y 2.8b-g 1.2j-n 8854.2i-p 1918.1n-v 44.6c-r 21.75m-a 11.110f 33.330e 33.330d

14 47g-j 119.5c-l 114.8a-g 4.1c-i 16.9c-i 3.2abc 2.1c-i 10294.1d-j 2098.7i-o 29.9z-b 20.39q-a 77.780a 55.560c 11.110e

15 47g-j 114h-m 90.1l-y 2.9mn 17.2c-h 2.5d-g 1.6f-m 8250l-t 1205.9j 28.2ab 14.625ab 55.560b 55.560c 55.560b

90
TRT DF DM PH PL NPP NSP NBP BY GY HSW HI AB(%) CS (%) RT (%)

16 52.5bc 118c-l 98.1f-u 3.4g-n 12.5h-t 2.5d-g 3.1a 10641c-h 1533.1b-g 30.35y-b 14.425ab 33.330d 33.330e 11.110e

17 47.5f-i 117.5c-m 107.4b-m 3.09l-n 10.8k-y 2.8b-g 1.4h-n 8370.5l-s 1931m-u 35.7p-b 23.135l-z 11.110f 33.330e 33.330d

18 46h-l 121b-j 112a-i 3.95c-l 7v-z 3a-e 1.3i-n 8411.7l-r 2239.4h-k 43.05c-t 26.755i-r 33.330d 55.560c 66.670a

19 44k-n 114.5g-m 93.9g-x 3.55e-n 7.8s-z 2.7b-g 1.1k-n 8631j-q 1477.6d-h 41.35d-w 17.105x-b 11.110f 33.330e 11.110e

20 44k-n 116e-m 104c-p 3.25i-n 9.4o-y 2.9a-f 1.4h-n 6801.5r-b 1972.1m-s 32.85s-b 29.245g-l 55.560b 55.560c 33.330d

21 46h-l 114h-m 114.5a-h 3.45g-n 13.9f-o 3.1a-d 1.8e-l 6798.2r-b 2222.4h-l 44.75c-q 34.575d-h 55.560b 11.110g 11.110e

22 44.5j-n 120b-l 101.4c-s 3.4g-n 9.4o-y 3.1a-d 2.1c-i 5572.8z-c 1926.2m-u 36.85m-a 34.755c-h 55.560b 33.330e 44.445c

23 51b-e 123a-g 99f-u 3.95c-l 8.6p-z 2.8b-g 3ab 7500o-w 2121i-n 42.95c-t 28.535g-n 33.330d 33.330e 11.110e

24 48.5e-h 119c-l 105.6c-n 4.1c-i 11.7i-w 3a-e 2.5a-e 8578.4k-q 2490.4e-g 50.4b-f 29.065g-m 33.330d 11.110g 11.110e

25 48f-i 119c-l 92.3i-y 4.05c-j 14.9e-n 3a-e 2.3a-g 9325.4g-n 2365.4f-h 36.35n-b 25.395j-u 33.330d 33.330e 11.110e

26 45.5i-m 114h-m 88m-y 3.65e-n 11.8i-w 2.4e-g 2.7a-d 6801.5r-b 1680.4w-d 39.7f-z 24.83j-w 33.330d 33.330e 33.330d

27 46h-l 118.5c-l 92.3i-y 3.65e-n 6.4x-z 2.7b-g 2.3a-g 5277.8bc 1383.1e-j 39.1h-z 26.265j-t 55.560b 33.330e 33.330d

28 42n 111.5lm 77v-y 3.2j-n 7v-z 2.7b-g 1.4h-n 7333.3p-x 1239.6ij 30.65w-b 16.92y-b 33.330d 55.560c 33.330d

29 46.5g-k 115g-m 74.2xy 3.3h-n 7.1v-z 2.5d-g 1.3i-n 2902d 1351.6f-j 33.95r-b 47.635a 33.330d 44.445d 66.670a

30 50c-f 116e-m 83q-y 3.35g-n 3.8z 2.4e-g 1.4h-n 7524.7o-w 1420.5e-j 42.3d-u 18.9t-b 33.330d 55.560c 55.560b

31 44.5j-n 114h-m 79.2u-y 3.4g-n 7.7s-z 2.2g 1.4h-n 5902.8w-c 1429e-j 39.4g-z 24.225k-y 22.220e 33.330e 33.330d

91
TRT DF DM PH PL NPP NSP NBP BY GY HSW HI AB(%) CS (%) RT (%)

32 46h-l 112k-m 102.3c-r 3.55e-n 15.7d-l 2.9a-f 1.9d-k 6761.7r-b 2156.2h-m 36.3n-b 31.935d-j 33.330d 66.670b 11.110e

33 44.5j-n 116e-m 99.2e-u 3.7e-m 13.2g-r 2.6c-g 1.3i-n 4861.1c 2320.3g-i 36o-b 48.655a 55.560b 33.330e 55.560b

34 46.5g-k 116.5d-m 93i-x 3.7e-m 12h-w 2.9a-f 1.7e-m 7352.9p-x 1824q-y 41.2e-x 24.81j-w 33.330d 33.330e 11.110e

35 43.5l-n 119.5c-l 90.6j-y 3.75e-m 7.2u-z 2.7b-g 1.2j-n 5468.8a-c 1507.1b-h 38.9h-a 28.095g-o 33.330d 33.330e 55.560b

36 45.5i-m 114.5g-m 111.3a-k 3.9d-l 21.7bc 2.8b-g 2.8abc 7623.8o-v 2610e 38.25j-a 34.26d-h 11.110f 55.560c 11.110e

37 42n 117d-m 105.5c-n 3.35g-n 12.5h-t 2.4e-g 2.1c-i 6561.1t-b 1798.4r-a 35.25p-b 27.72h-q 33.330d 55.560c 11.110e

38 42.5n 116.5d-m 97.5f-v 4.35c-f 13.8g-p 2.9a-f 1.8e-l 7843.1n-u 1861.7p-x 40.6e-z 23.795k-y 11.110f 33.330e 11.110e

39 44.5j-n 112k-m 92.6i-x 3.15k-n 7.3t-z 2.4e-g 2.1c-i 5915.2w-c 1590z-e 32.9s-b 27.99g-p 55.560b 55.560c 33.330d

40 53.5b 121b-j 89.5l-y 3.7e-m 11.3j-x 3a-e 1.7e-m 7572.1o-w 1538.5b-g 38.6i-a 21.39n-a 33.330d 44.445d 44.445c

41 44k-n 122.5a-h 103.5c-q 4.1c-i 19.1b-f 2.9a-f 1.8e-l 14186.3a 3422.3b 46.8b-n 24.145k-y 33.330d 11.110g 11.110e

42 46h-l 116e-m 120.3a-d 4c-k 22.8b 2.8b-g 3.1a 8051.5m-u 3582.7b 35.1q-b 44.655ab 33.330d 33.330e 11.110e

43 48f-i 119c-l 120.1a-e 3.6e-n 19.8b-e 2.8b-g 2.2b-h 11937.5bcd 3528.7b 43.15c-t 29.59g-l 11.110f 11.110g 11.110e

44 42n 115g-m 84.3o-y 3.2j-n 12.2h-v 2.7b-g 1l-n 12192.7bc 1773.1s-a 50.3b-f 14.54ab 33.330d 33.330e 11.110e

45 43mn 115g--m 79.5t-y 3.55e-n 7.1v-z 2.85a-g 1.6f-m 6806.7r-b 1323.9g-j 25.8b 19.495r-b 77.780a 77.780a 55.560b

46 44.5j-n 118.5c-l 103c-r 4.1c-i 7.3t-z 2.8b-g 1.8e-l 9843.8e-l 1733.4t-b 48.15b-l 17.605w-b 33.330d 11.110g 11.110e

47 43mn 116.5d-m 96.3f-w 3.45g-n 10.5l-y 2.6c-g 1.5g-m 6125v-c 1956.6m-t 45.75b-q 31.97d-j 55.560b 22.220f 55.560b

92
TRT DF DM PH PL NPP NSP NBP BY GY HSW HI AB(%) CS (%) RT (%)

48 42n 114h-m 97f-v 3.2j-n 9o-z 2.5d-g 1.2j-n 8893.4i-p 1711.6u-c 35.3p-b 19.27s-b 33.330d 33.330e 11.110e

49 42.5n 109m 109.1b-l 3.4g-n 18.4b-g 2.7b-g 1.9d-k 9625f-m 2941d 38.75i-a 30.635e-k 11.110f 11.110g 11.110e

50 43.5l-n 115g-m 93i-x 3.35g-n 7.5t-z 2.4e-g 1.1k-n 5777.3x-c 1600.5y-e 37.3m-a 29.615g-l 33.330d 11.110g 33.330d

51 42n 118c-l 85n-y 3.35g-n 6.8w-z 2.4e-g 1.6f-m 5902.8w-c 2222.3h-l 41.25e-x 37.8b-e 55.560b 33.330e 55.560b

52 47.5f-i 128.5ab 120.5abc 4.4cde 14.8e-n 2.7b-g 2c-j 8072.9m-u 2003.3l-s 53.3bc 24.835j-w 33.330d 11.110g 11.110e

53 44k-n 121.5a-i 90l-y 3.55e-n 9.7n-y 2.8b-g 1.1k-n 7582.7o-w 1868.6o-x 48.4b-k 24.665j-w 33.330d 33.330e 33.330d

54 44k-n 117.5c-m 90.5k-y 3.75e-m 8.9o-z 2.8b-g 1.7e-m 7009.8q-a 1468.3d-i 42.3d-u 20.965o-a 33.330d 66.670b 11.110e

55 42.5n 117d-m 97.7f-v 3.85e-l 10.7k-y 2.6c-g 1.3i-n 6066.2v-c 2042.7k-q 47.35b-m 33.71d-i 55.560b 33.330e 33.330d

56 46.5g-k 120.5b-k 91j-y 3.85e-l 11.3j-x 3a-e 1.5g-m 5416.7a-c 1817.8q-z 37.4l-a 35.29c-g 55.560b 33.330e 33.330d

57 44k-n 120b-l 82.1r-y 3.85e-l 8.2q-z 3.1a-d 1.3i-n 7239.6p-z 1916n-v 31.8u-b 26.545i-s 55.560b 55.560c 11.110e

58 47.5f-i 124.5a-e 106.5b-m 3.35g-n 15.3d-m 2.9a-f 1.3i-n 10959.6c-g 1928.7m-u 30.55x-b 17.625w-b 33.330d 11.110g 11.110e

59 43mn 121.5a-i 98.1f-u 3.6e-n 18.3b-g 3a-e 1.5g-m 9161.2h-o 3141.8cd 45.95b-p 34.305d-h 33.330d 33.330e 11.110e

60 44.5j-n 115.5f-m 87m-y 3.3h-n 10.7k-y 2.4e-g 0.9mn 6454.3u-c 1927.6m-u 43.2c-s 30.29g-l 33.330d 55.560c 33.330d

61 42n 117.5c-m 96f-w 3.4g-n 13.4g-q 3.1a-d 1l-n 4843.8c 2034k-q 41.9d-v 42abc 33.330d 33.330e 66.670a

62 44k-n 112k-m 126.7ab 4.1c-i 36.9a 3.3ab 2.8abc 13125ab 5004.4a 38j-a 38.155bcd 11.110f 11.110g 11.110e

63 50c-f 121.5a-i 116a-f 5.8a 13.2g-r 3.1a-d 2.4a-f 12111.1bc 3011.7cd 80.1a 25.035j-v 11.110f 11.110g 11.110e

93
TRT DF DM PH PL NPP NSP NBP BY GY HSW HI AB(%) CS (%) RT (%)

64 42.5n 118c-l 102c-r 3.75e-m 19.9b-e 2.8b-g 2.1c-i 14276a 3511.8b 43.95c-r 24.6j-w 33.330d 11.110g 11.110e

65 43.5l-n 118.5c-l 93.5i-x 3.65e-n 10.8k-y 2.9a-f 1.7e-m 8137.3m-u 2356.8f-h 41.05e-y 28.97g-m 33.330d 33.330e 11.110e

66 44k-n 119c-l 99.5d-u 3.9d-l 9.5o-y 2.8b-g 1.4h-n 9194.4h-o 1831.1q-y 50.7b-e 19.955r-b 33.330d 33.330e 11.110e

67 44k-n 120.5b-k 93i-n 3.95c-l 7.9s-z 3.2abc 1.7e-m 8114m-u 2074.6j-p 50.45b-f 25.575j-u 11.110f 33.330e 11.110e

68 43.5l-n 123a-g 97.3f-v 3.6e-n 8.9o-z 3a-e 1l-n 7582.7o-w 1904.9n-w 38.05j-a 25.135j-v 11.110f 33.330e 11.110e

69 42.5n 122a-h 100c-u 4.75bcd 6.9w-z 2.8b-g 1.2j-n 8631j-q 2583.8ef 56.3b 29.97g-l 11.110f 33.330e 11.110e

70 44k-n 118.5c-l 98.8f-u 3.5f-n 16.1d-j 2.8b-g 1.5g-m 8107.8m-u 3053.7cd 41.1e-y 37.705b-f 33.330d 11.110g 33.330d

71 43.5l-n 121.5a-i 102.1c-r 4.15c-h 11.3j-x 2.9a-f 1.5g-m 8864i-p 1823.8q-y 44.15c-r 20.605p-a 33.330d 11.110g 11.110e

72 44k-n 120b-l 99.4d-u 4.8bc 9.8n-y 2.7b-g 1.3i-n 8631j-q 1847.9p-x 49.6b-h 21.385n-a 11.110f 11.110g 33.330d

73 43.5l-n 114h-m 87.5m-y 3.15k-n 6y-z 2.5d-g 1.4h-n 7287.3p-y 1427.4e-j 44.05c-r 19.625r-b 55.560b 55.560c 33.330d

74 43.5l-n 121b-j 111.2a-k 3.95c-l 11.7i-w 2.7b-g 1l-n 13111.1ab 3479.5b 48.7b-j 26.62i-s 33.330d 11.110g 11.110e

75 42.5n 114h-m 94.2g-x 3.3h-n 7.6s-z 2.7b-g 1.1k-n 7290.9p-y 1415.4e-j 38.45i-a 19.455r-b 55.560b 33.330e 33.330d

76 42n 120.5b-k 98f-u 3.9d-l 7.9s-z 2.9a-f 1l-n 7415.2p-x 1570.7a-f 49.2b-i 21.18n-a 33.330d 33.330e 33.330d

77 44k-n 126a-c 100.5c-s 3.45g-n 12.4h-u 2.7b-g 1.1k-n 9361.1g-n 1928.3m-u 46.55b-o 20.595p-a 55.560b 33.330e 11.110e

78 43.5l-n 130a 127ab 6.15a 20.5bcd 3.2abc 2.1c-i 14805.6a 2694.8e 88.75a 18.205u-b 11.110f 11.110g 11.110e

79 46h-l 122.5a-h 96f-w 4.1c-i 11.5j-x 2.9a-f 1.6f-m 8117.6m-u 1883.3o-x 44.1c-r 23.305k-z 33.330d 33.330e 33.330d

94
TRT DF DM PH PL NPP NSP NBP BY GY HSW HI AB(%) CS (%) RT (%)

80 42n 120b-l 111.5a-j 4.15c-h 12h-w 2.6c-g 1.1k-n 7037.8q-a 2102.2i-o 52.05bcd 30.345f-l 11.110f 33.330e 11.110e

81 42n 122a-h 131.5a 5.4ab 15.9d-k 2.9a-f 2.7a-d 13131.9ab 3179.8c 78.1a 24.325k-x 55.560b 11.110g 11.110e
Note: TRT= treatment, DF= days to flowering, DM= days to maturity, PH= plant height, PL= pod length, NPP= number of pods per plant,
NSP=number of seeds per pod, NBP= number of branches per plant, BY= biomass yield, GY= grain yield, HSW= 100 seed weight, HI=
harvest index, AB= ascochyta blight, CS= chocolate spot, RT= rust disease

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 BIOGRAPHICAL SKETCH

The author, Andualem Muche, was born on 12 June 1996 in Mecha district West Gojjam to
his father Muche Hiywotu and his mother Gedam Addis. When he reached school age, he
enrolled at Kudmi Elementary and Junior School at Kudmi to attend elementary and junior
education. In 2011 he then joined Merawi high School in Merawi to attend his secondary and
preparatory education. After accomplished his high school in 2015, he joined Gambella
University and graduated with the Degree of Bachelor of Sciences in plant science in 2018.
Upon graduation, he joined the School of Graduate Studies at Bahir Dar University in 2018 to
pursue a study leading to the Degree of Master of Science in plant breeding.

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