Besufekad (1)
Besufekad (1)
Besufekad (1)
M.Sc. THESIS
BESUFEKAD ENIDEG
JANUARY, 2012
JIMMA UNIVERSITY
GENETIC VARIABILITY AND CHARACTER ASSOCIATION IN
SOYBEAN (Glycine Max L. Merrill) GENOTYPES
BY
BESUFEKAD ENIDEG
JANUARY, 2012
JIMMA UNIVERSITY
SCHOOL OF GRADUATE STUDIES
JIMMA UNIVERSITY
APPROVAL SHEET
As Thesis Research advisor, I hereby certify that I have read and evaluated this thesis prepared,
As member of the Board of Examiners of the M.Sc. Thesis Open Defense Examination, We
certify that we have read, evaluated the Thesis prepared by Besufekad Enideg and examined the
candidate. We recommended that the Thesis be accepted as fulfilling the Thesis requirement for
i
DEDICATION
ii
STATEMENT OF THE AUTHOR
First, I declare that this thesis is my original work and that all sources of materials used for this
thesis have been dully acknowledged. This thesis has been submitted in partial fulfillment of the
requirements for M. Sc. degree in Plant Breeding at the Jimma University and is deposited at the
university Library to be made available to borrowers under rules of the library. I solemnly declare
that this thesis is not submitted to any other institutions anywhere for award of any academic
Brief quotations from this thesis are allowable without special permission provided that accurate
acknowledgment of the source is made. Requests for permission for extended quotation from or
reproduction of this manuscript in whole or part may be granted by the Head of the Department of
Horticulture and Plant Sciences or the Dean of Graduate Studies when in his judgment the
proposed use of the material is in the interest of scholarship. In all other instances, however,
iii
BIOGRAPHICAL SKETCH
The author was born on July 22, 1985 in Addis Ababa, Ethiopia. He attended his primary
school at Elay Elementary school from 1993 to 1998, and RasGobena elementary school from
1999 to 2000, and secondary high school at Gambella Senior Secondary School from 2001 to
2004.
The author joined Haramaya University in 2005 and completed his undergraduate studies with
B.Sc. degree in Plant Sciences in July 2007. He was employed at Gambella Agricultural Research
Institute starting from September 2007 as a Junior Researcher until he joined Jimma University in
September 2009 as a postgraduate student with the specialization in plant breeding through the
sponsorship of Rural Capacity Building Project (RCBP).
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ACKNOWLEDGMENTS
Praise be to God, the most gracious and the most merciful who has made everything happen, who
is the entire source of all knowledge and kinds of wisdom endowed to mankind. My deep and
sincere gratitude goes first to my major advisor Dr. Sentayehu Alamerew for his kind, precise
and keen interest as well as generous thoughts and encouragements offered to me throughout
the thesis research work. I am deeply indebted to my Co-advisor Mr. Abush Tesfaye for his
critical evaluation of the thesis research work in the field at Melko (Jimma), providing me the
tested materials, technical guidance made during data organization and biometrical analysis as
well as his constructive comments were quite useful and are gratefully acknowledged.
I am forever grateful to my dearest friend Maria Liu Wong and her friends who bought me a
laptop computer from New York, without that this thesis work would have not been
completed in time and for her unrestricted support at each turning point of the road. I will
never forget.
I am also deeply indebted to the staff of Jimma Agricultural Research Center (JARC)
especially the management of the center, AtoBehailuAtero and BerhanuBekele for their
unreserved support throughout the thesis work at the field and for allowing me to use the
laboratory facilities.
My special gratitude extends to my dearest friends AtoTamratKassa for his support during the
thesis write up, Miss HarnetAbrha and Miss SerkalemGetahun for helping me during the data
processing.
Finally, those of you, who helped me in many ways and whose names are not mentioned here,
I thank you very much.
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ABBREVIATIONS
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TABLE OF CONTENTS
Contents Page
vii
TABLE OF CONTENTS CONTINUED
viii
LIST OF TABLES
Tables Page
ix
LIST OF TABLES IN THE APPENDICES
I. Analysis of variance for mean square of the 15 characters of49 soybean genotypes tested at
Jimma (2010/2011) ............................................................................................................ 83
II. Analysis of variance for mean square of the 15 characters of 49 soybean genotypes tested
at Assosa (2010/2011) ........................................................................................................ 84
III. Analysis of variance (ANOVA) for the combined characters of 49 soybean genotypes
tested at Jimma and Assosa (2010/2011) ........................................................................... 85
IV. Mean values of 15 traits of soybean genotypes grown at Jimma (2010/11) ...................... 86
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LIST OF FIGURES
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GENETIC VARIABILITY AND CHARACTER ASSOCIATION IN SOYBEAN (Glycine
max L. Merril) GENOTYPES
ABSTRACT
Information on the extent and pattern of genetic variability, interrelationship among different
agronomic characters and knowledge of diversity are essential to design breeding strategies
in the available germplasm of soybean and helps to identify elite genotypes that will be
incorporated into soybean crop improvement programs to address the growing demand of the
crop in Ethiopia. Forty-nine soybean (Glycine max (L.) Merrill) genotypes were tested in 7x7
simple lattice design at Jimma and Assosa with the objectives of estimating genetic variability
and associations among characters, and to estimate genetic divergence and, thereby, to
cluster the test genotypes into genetically divergent classes. Analysis of variance revealed that
there was statistically significant difference among the forty nine genotypes for most of the
traits studied except root volume and root dry weight at Jimma. The relatively wide range of
the mean values for most of the characters indicated the existence of variations among the
tested genotypes. High phenotypic coefficient of variation (PCV) and genotypic coefficient of
variation (GCV) was recorded for grain yield, biomass yield, pod number per plant, plant
height, total nodules per plant, effective nodules per plant, and harvest index at both
locations. The highest heritability value was recorded for grain yield at both locations. High
heritability, coupled with high expected genetic advance as percent of mean, was observed for
grain yield, harvest index, biomass yield, total nodules per plant, effective nodules per plant
and pod number per plant across both locations. This indicates that the characters can be
improved through selection. Days to 50% flowering, days to pod setting and days to maturity
showed negative and significant genotypic and phenotypic association with grain yield at
Jimma. Grain yield was negatively and significantly correlated with biomass yield, pod
number per plant and hundred seed weight both at genotypic and phenotypic levels at Assosa.
Genotypic path analysis revealed that effective nodules per plant and pod number per plant at
Jimma and Assosa, respectively exerted the highest positive direct effect. The D2 analysis
grouped the 49 soybean genotypes into five and three distinct clusters at Jimma and Assosa,
respectively. The principal component analysis revealed that 6 and 5 principal components at
Jimma and Assosa, respectively have accounted for 79.90% and 73.81% of the total variation,
respectively.
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1. INTRODUCTION
Soybean Glycine max (L.) Merrill] (2n= 14) belongs to the family Leguminacae, subfamily
Papilionoideae, tribe phaseolae, and genus Glycine. It is reported to be originated in Asia,
probably in north eastern China about 2500 B.C. (Poehlman and Sleeper, 1995). Since then, it
has spread to different countries in the world and become an established component of world
agriculture.
It is the world’s leading source of oil and protein. It has the highest protein content (40%) of
all food crops and is second only to groundnut in terms of oil content (20%) among food
legumes (Norman et al., 1995; Soy Stats 2008; Poehlman and Sleeper, 1995). The meal is also
rich in minerals, particularly calcium, phosphorus and iron (Ogokeet al., 2003).
The majority of the soybean crop is processed into oil and meal. Oil extracted from soybeans
is made into shortening, margarine, cooking oil, and salad dressings. Soybeans account for 80
percent or more of the edible fats and oils consumed in the United States. Soy oil is also used
in industrial paint, varnishes, caulking compounds, linoleum, printing inks, and other products.
Development efforts in recent years have resulted in several soy oil-based lubricant and fuel
products that replace non-renewable petroleum products (Gibson and Benson, 2005).
Since soybean is a leguminous crop, it fixes its own nitrogen in association with
Bradyrhizobiurnjaponicum. If soybeans have not been grown on the field or it has been many
years since soybeans were raised, an inoculant should be applied at planting to establish the
bacteria in the soil (Hoeftet al., 2000).
The soybean is a day length sensitive crop. Length of daylight is the principal factor that
affects the amount of vegetative growth before flowering begins. The ideal situation is that the
plants grow to a reasonable size (2-3 feet) before they bloom. Large plants tend to bear a large
number of seeds. Thus, seed yield potential per plant is closely related to the day length
requirement of the variety and to the season of planting. It is recommended, therefore, that in
the preliminary stages of developing soybean as a crop in a new region, several varieties be
tried as well as several planting dates, and that careful notes be taken including planting date,
date of flowering, harvest date, and number of seeds per plant (Gibson and Benson, 2005).
Soybean is a hot weather crop suitable for year-round growth in most parts of the tropics.
Temperatures of at least 150 c are needed to germinate the seed and mean temperatures of 20-
25 0c to grow the crop. Soybeans need at least moderate soil moisture in order to germinate
and for seedlings to become established, but need dry weather for the production of dry seed.
Soybeans suffer if the soil is waterlogged.
In 2007, the total cultivated area of soybean in the world was 90.19 million hectares and the
total production was 220.5 million tons (FAO, 2009).United States of America is the leading
soybean producer and exporter with annual production of 70 million metric tonnes in
2007.The second largest producer and exporter Brazil produced 61.0 million metric tonnes in
the same year. Argentina, China and India produced 47, 14.3 and 9.3 million metric tons
respectively in the year 2007 (USDA, 2008).
In 2005, the U.S. was the number one soybean consumer in the world with a total amount of
51 million tones followed by Brazil 32 million tones, Argentina 31 million tones and China
25 million tons (USDA, 2008). In 2007/08 a total of 236.8 million tons of soybean was
produced worldwide (FAO, 2008).
Twenty-one African countries now produce soybean. Nigeria has the highest 6-year (2000-05)
average production of 486,000 tons on an area of 553,260 hectares, followed by South Africa
with 205,270 tons from 122,870 hectares, and Uganda with 155,500 tons from 139,500
hectares (IITA 2008).
In Ethiopia, soybean is grown over wider agro-ecologies especially in low to mid altitude
areas (1300 to 1700 masl) that have moderate annual rainfall (500-1500mm) (Gurmuet al,.
2009). The area covered under soybean is 6,236 hectares and the total production of the crop
in the country is 78,989 quintals. The productivity of the crop is 12.67 quintals per hectare
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(CSA, 2008) which is very low as compared to the productivity of the crop in the world
(FAO, 2009). This is attributed to lack of improved varieties, biotic factors such as diseases
and insect pests, and abiotic factors.
Diseases such as rust, red leaf blotch, frog-eye leaf spot, bacterial pustule, bacterial blight, and
soybean mosaic virus are problems to be resolved in soybean. Soybean rust
(Phakopsorapachyrhizi) particularly is the most destructive foliar disease of soybean in recent
times, and can cause 50–60% yield loss. It is a major disease worldwide. Among insect pests,
pod sucking and defoliating insects are major constraints (IITA, 2009).
Lack of varieties tolerant to midseason moisture stress and high yielding varieties tolerant to
low phosphorus are among the abiotic constraints. Research on seed quality such as protein,
oil, carbohydrate, and anti-nutritional factors is lacking. Moreover, lack of emphasis on using
molecular markers as aid to conventional breeding is also worth mentioning (IITA, 2009).
Much effort has been made to improve soybean productivity in Ethiopia since conception of
soybean breeding in the country (Asfawet al. 2003). As a result some varieties have been
released. For further improvement of the crop the knowledge of variability and association of
yield and its related traits is essential. Therefore this research project was conducted with the
following objectives:
To estimate the extent of phenotypic and genotypic variability, heritability and the
genetic advance expected under selection.
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2. LITERATURE REVIEW
2.1.1. Taxonomy
According to recent taxonomical classification, soybean belongs to the genus Glycine, which
has two subgenera: Soja and Glycine. Cultivated soybean (G. max) and its wild annual
relative G. soja belong to the subgenus Soja. The subgenus Glycine contains 16 wild
perennial species, mostly found in Australia. All of these species generally carry 2n = 40
chromosomes, except for G. hirticaulis, G. tabacina and G. tomentella (Vaughan and
Hymowitz, 1983; Brown et al., 1987; Hymowitzet al., 1997). Some of these wild perennial
species also have polyploidcytotypes. Glycine is believed to be an ancient polyploid having ×
= 10; however, plants with 2n = 40 behave cytologically like diploids. The annual Glycine is
derived from the perennial forms.
The subgenus Soja is most diverse in the eastern half of north China, whereas maximum
diversity for the subgenus Glycine occurs in Australia. The wild perennial Glycine species
found outside of Australia were taken to other neighboring regions by migratory birds via
long distance dispersal (Hymowitzet al., 1997).
2.1.2. Evolution
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The genus Glycine is thought to be of ancient polyploid origin due to the high chromosome
number of the majority of the species (n = 20) compared to closely related genera (mostly n =
10 or 11, one with n = 14; Goldblatt, 1981). Additional lines of evidence exist, including
cytogenetic studies in haploid G. max (Crane et al., 1982), supporting this hypothesis of
polyploidy origin. Schuelteret al. (2004), found that the Glycine genome has gone through
two major rounds of duplication, the first estimated at 41.6 million years ago and another at
14.5 million years ago. Van et al. (2008), looked at evolutionary events, revealing that the
recent divergence of two soybean homologous regions occurred at 60 and 12 million years
ago, respectively. Clarindoet al. (2007) found that the karyograms support soybean’s
tetraploid nature (4× = 40), specifically for the presence of chromosomes with identical
morphology, and suggested that chromosome rearrangements may have occurred during the
speciation of G. max.
The genus Glycine Willd.is divided into two subgenera, Glycine (perennials) and
Soja(Moench) F.J. Herm. (annuals). The perennial species are extremely diverse in
morphology, cytology and genome composition. They grow in very diverse climatic and soil
conditions and have a wide geographic distribution. The species have been screened for many
physiological and biochemical traits as well as for sources of resistance to economic
pathogens. Some perennial Glycine species are sources of resistance to soybean cyst
nematode and a source of lack of Bowman-Birk protease inhibitor (Hymowitz, 2004).
2.1.3. Distribution
Soybean is believed to be of Chinese origin, having been derived from a slender, twig-like
plant known as G. ussuriensis. Nagata (1960), suggested that the species originated in China
proper, probably in the north and central regions. Piper and Morse (1923), considered that the
wild form G. ussuriensis was known to occur in China, Manchuria and Korea and stated that
soybean is native of eastern Asia. According to Hymowitz (1970), G. ussuriensis grows wild
in Korea, Taiwan and Japan throughout the Yangtze valley, the northern provinces of China
and the adjacent areas of the former USSR. Based on cytogenetic evidence, Hymowitz (1970)
concluded that G. max and G. ussuriensis are the same species and also stated that historical
5
and geographical evidence points to the eastern half of northern China as the area where
soybean was first domesticated around the 11th century BC. Nagata (1960) suggested that the
cultivated form of soybean was introduced into Korea from China and then disseminated to
Japan between 200 BC and the third century AD. Shipments of soybeans and soybean
products were made to Europe around 1908 and soybean attracted worldwide attention. Aiton
(1814) indicated that soybean was first brought to England in 1790 and cultivated at the Royal
Botanic Gardens, Kew, in that year.
Hymowitz and Barnard (1991) made a detailed account of early introductions in the USA and
mentioned that during the first two decades, new soybean accessions were introduced from
India and China into the USA by plant explorers.
Soybean was introduced to neighboring countries (Japan, India, Nepal, Russia) from China
around the first century AD. It appears that missionaries may have been the first to bring
soybean to Europe early in the 18th century. Soybean was first introduced to the USA in 1765
(Hymowitz and Harlan, 1983) and was then spread to Canada and Latin America. Soybean
production began only recently in Africa, during the second half of the 20th century. Soybeans
were taken to Brazil and Argentina in 1822 and 1862, respectively (Larreche and Brenta,
1999).
Thousands of soybean landraces with great genetic diversity have been selected and preserved
by Chinese farmers during a long history of cultivation. The Yellow River region of China is
generally considered as the centre of origin of soybean, based on the existence of a great
number of wild soybeans and the earliest record of soybean in China (Hymowitz and
Kaizuma, 1981). Wild soybean (G. sojaSieb. andZucc.) is widely distributed in nearly all
provinces of China, Korea, Japan and parts of Russia (Hymowitz and Singh, 1987). Based on
tremendous diversity in cultivated and wild soybean, China has collected 23,000 accessions of
G. max. In addition, 5300 accessions of G. soja have been conserved in a gene bank for long-
term storage.
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Thirteen wild perennial species of soybean collected by USDA explorers are indigenous to
Australia (Hymowitz and Bernard, 1991). All carry 2n = 40 chromosomes. G. tabacina
(Labill.)Benth., with 2n = 40 or 80 chromosomes, has been found in Australia, Taiwan, the
South Pacific Islands and the islands of the west central Pacific. All accessions of G. tabacina
collected outside of Australia are tetraploid (2n = 80) and, even including Australia, the
tetraploid predominates over the diploid form (Singh et al., 1987, 1989; Hymowitz and
Bernard, 1991), demonstrating that the complexes of G. tabacina and G. tomentella evolved
through alloploidy in Australia. This clearly indicates that the wild perennial species of
soybean have invaded Australia and associated areas, and the wild annual G. soja has invaded
central and northern Asia. Since G. soja is the wild ancestor of soybean (Hymowitz and
Newell, 1981) and all morphological and genetic variability exist in China in the form of
landraces and primitive cultivars, this indicates that China is the centre of diversity.
Developing crop cultivars with high seed and oil yield and with desirable nutritional and feed
quality has been the principal aim of soybean breeding programs worldwide. The strategy of
crop improvement of any trait comprises the collection or generation of highly ranking variant
types of populations and the progressive reduction of them by selection (Ashley, 1999).
The presence of variation in the germplasm for the trait of interest is, therefore, very
important. 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).
7
Genetic variability, which is due to the genetic differences among individuals within a
population, is the core of plant breeding because proper management of diversity can produce
permanent gain in the performance of plant and can buffer against seasonal fluctuations
(Welsh, 1990; Sharma, 1998). In addition, estimation of the magnitude of variation within
germplasm collections for important plant attributes will enable breeders to exploit genetic
diversity more efficiently (Jahufer and Gawler, 2000).
As in other major crops, genetic diversity of soybean commercially grown cultivars has been
decreasing at an alarming rate. The narrowness of North American (Gizliceet al., 1994;
Sneller, 1994) as well as Brazilian (Velloet al., 1984) soybean germplasm has been well
documented by pedigree analyses. Gizliceet al. (1994) determined that only 35 ancestors
contributed more than 95 % of all alleles and only five lines account for more than 55 % of
the genetic background of public cultivars in North America. Similarly, Gaiet al. (1998) and
Wang et al.,(2008), reported that 651 soybean cultivars released from 1923 to 1995 in China
could be traced back to only 308 ancestors or about 1.5% of the germplasm resources
available in China.
According to Poehlman (1979) and Welsh (1981), dissimilarity will always exist among
individuals in a population and assessing the origin and magnitude of variability is the key to
success in a crop improvement program. Frey (1981) indicated that the extent of the genetic
variability in a specific breeding population depends on the germplasm included in it. Hence,
genetic variability is of immense importance to plant breeders because it can be transmitted to
the progeny and the proper management of the diversity can produce permanent gain in the
performance of the plant (Welsh, 1981).
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Oyaet al. (2004) in the study for drought tolerance characteristics of 11 Brazilian soybean
cultivars, wide range of variability was observed on the basis of seed yield among all the
cultivars studied. The report also indicated that in cultivars with higher drought tolerance crop
growth rate during the drought stress period was higher than in other drought susceptible
cultivars.
Hufstetleret al., (2007) studied genetic variability in 23 cultivars of soybean for the three
physiological traits that may affect performance of soybean when soil water availability is
limiting viz. water use efficiency (WUE), regulation of whole plant water use in response to
soil water content, and leaf epidermal conductance (ge) when stomata are closed. They
reported significant variation (P<.001) among the genotypes for the three traits that determine
drought tolerance ability of the soybean crop. Significant differences were found among
genotypes for minimum ge values (P <.001) across leaf positions, but there was no difference
in ge on the basis of leaf position (upper vs. lower), and no genotype by leaf position
interactions were present.
Tahiret al. (2009), conducted a research to evaluate the effect of rhizobium inoculation and
NP fertilization on growth, yield and nodulation of soybean genotypes the result revealed a
wide range of variability among the rhizobium inoculation levels and treatment combinations
of NP fertilizer levels and demonstrated significant increase of grain yield and nodulation
compared to the un-inoculated genotypes.
2.3 Heritability
9
Heritabilitycanbedefined,inbroadsense,astheproportionofthegenotypicvariabilitytothetotalvari
ance(Allard,1960).Itreferstotheportionofphenotypicallyexpressed
variation,withinagivenenvironmentanditmeasuresthedegreetowhichatraitcanbe modified
byselection(Christianson andLewis , 1982). According
toFalconerandMackay(1996),heritabilityinnarrowsenseisdefinedas
“theratioofadditivegeneticvariancetophenotypicvariance”.Sincebroadsenseheritabilitydoesnot
giveaclear pictureoftransmissibilityofvariationfromgenerationtogeneration(becausethegenetic
variationincludesthefixableandnon-fixabledominanceandepistaticvariation),itsutilization
islimited inplant improvementprogram.
Incontrast,estimateofheritabilityinanarrowsensecangiveclearerpicturethanthatofbroadsense(Fa
lconerandMackay, 1996). Estimationofheritabilityasaratioofgenotypic
tophenotypicvariancemayvarygreatly dependingup on
theunitforwhichvarianceisconsidered(Johnsonetal.,1955a).
It is obvious that difference due to environment may tend to obscure genotypic variations.
The greater the proportion of the total variability that is due to the environment the more
difficult it will be to select for inherited differences. On the other hand, if environmental
variability is small in relation to heritable differences, selection will be efficient because the
characters to be selected will be transmitted to its progeny (Briggs and Knowles, 1987).
If genetic variation in a progeny is large in relation to the environmental variation the
heritability will be high or if genetic variation is small in relation to the environmental
variation, then heritability will be low (Mittal and Sethi., 2004).
High heritability was reported by Adityaet al, (2011), for three characters viz, days to 50 per
cent flowering, number of primary branches per plant and 100 seed weight (91%) in 31
soybean genotypes.
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., 1955a). 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.
11
Adityaet al, (2011), reported high heritability coupled with high expected genetic advance in
31 soybean genotypes for number of pods per plant and dry matter weight per plant.
Genetic progress would increase with increase in the variance. Therefore, the utility of
estimates of heritability is increased when they are used in conjunction with the selection
differential, the amount that the mean of the selected lines exceeds the mean of the entire
group (Johnson etal., 1955a). According to Burton and DeVane (1953), genetic advance tell
us the estimate of the expected gain for a particular character through selection. Shivakumar
(2008), in the study for genetic variability and character association in 64 soybean genotypes
reported high heritability coupled with high genetic advance as percent of mean for grain
yield and pod number per plant.
The various characteristics of crop plants are generally interrelated or correlated. Such
correlations can be either negative or positive. In plant breeding and genetic studies,
correlated characters are of prime importance because genetic causes of correlations through
pleiotropic action or developmental interactions of genes and changes brought about by a
natural or artificial selection (Singh, 1993; Falconer and Mackay, 1996; Sharma, 1998).
The relative influence of various traits and their degree of associations can be estimated
statistically by correlation (Dewey and Lu, 1959). Determination of relationships of characters
can help to identify traits of economic importance.
Generally, three types of correlations are discussed in quantitative genetics and these are
phenotypic, genotypic and environmental correlations. The association between two characters
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 characters are linearly
related. It is determined from measurements of the two characters 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 characters.
12
Studies on genotypic and phenotypic correlations among characters of crop plants are useful
in planning, evaluating and setting selection criteria for the desired characters in breeding
program (Johansson et al., 1955b). Correlations between different characters 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 characters.
Pleiotropy is the property of a gene, which affects two or more characters; as a result it causes
simultaneous variations in the two characters when the genes are segregating (Singh, 1993;
Falconer and Mackay, 1996).
Correlation coefficient analysis helps to determine the nature and degree of relationship
between any two measurable characters. Characters that are not easily measured or which are
largely influenced by the environment have low heritability ratio; hence, there is a need to
examine the relationships among various characters. Knowledge of the correlations that exist
between important characters may facilitate the interpretations of the results that are already
obtained, and provides the basis for planning more efficient breeding program. However, as
the number of independent variables influencing a particular dependent variable increases, a
certain amount of interdependence is expected. Therefore, correlation may be insufficient to
explain the associations in a manner that will enable one to decide on either a direct or an
indirect selection strategy (Bhatt, 1970).
Correlation coefficients may range in value from -1 to +1. Phenotypic correlations can
normally be estimated with a high degree of accuracy. Estimates of genetic correlations
however, usually have high standard errors because of difficulties to avoid the directional
effects of confounding factors (i.e. dominance and epistatic genetic effects) on additive
genetic correlation estimates (Lynch and Walsh, 1998 in Amsal 2001). In addition, genetic
13
correlations are strongly influenced by gene frequencies, and, therefore, may differ markedly
in different populations (Falconer and Mackay 1996).
Estimate of correlations and significance test was previously discussed by several workers
(Robertson 1959; Singh and Chaudhary 1977; Sharma, 1998). Depending on the sign genetic
correlations between two characters can either facilitate or impede selection progress.
Correlation value (r = 1) implies perfect (100%) correlation, where both traits vary hand in
hand, r = -1 means there is 100 % correlation between two characters, but they vary in
opposite direction, and r = 0 carries the implication that there is no correlation at all between
the two characters (Falconer and Mackay 1996; Sharma 1998).
Correlated characters 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 character with its fitness, which is the primary
agent, that determines the genetic properties of that character 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 character as a means of
improving another depends on the extent to which improvement in major characters is
facilitated by selection for the indicators. Such improvement depends not only on the
genotypic correlation but also phenotypic correlation (Johnson et al., 1955b).
For selection based on yield component to be effective in increasing yield, Sidwellet al. (1976)
stated that the components should fulfill the following: they should be highly heritable, the
component should be genotypically independent or genotypic correlation among the
component should be positive and the component should be physiologically related in a
positive manner.
14
Character association 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 character into some agronomically
superior cultivars from wild or uneconomic genotypes (Saraf and Hedge, 1984). Seed yield is
a polygenically controlled complex character 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).
Vasicet al. (1997) found correlations of plant height and productive height with yield, which
were established, via the number of pods per plant and the number of seeds per plant. These
results give a clear indication that the yield components are mutually very closely associated.
Thus, they concluded that productivity was more dependent on the number of pods per plant
than on the number of seeds per pod because the latter characteristic was quite stable in the
climatic region. The authors exhibited a positive direct correlation between seed size and
yield, which was masked by the negative correlation between seed size and the number of
pods per plant. 100-seed weight is positively and strongly correlated with seed length and
seed height (Zevenet al., 1999) but negatively correlated with number of pods per plant
(Nienhuis and Singh, 1986). Seed length is positively correlated with pod length and seed
height (Zevenet al., 1999). They also found positive correlations between pod length and
number of seeds/pod, 100-seed weight, seed length and seed height.
Although correlation estimates are helpful in determining the components of complex trait
such as yield, they do not provide an exact picture of the relative importance of direct and
indirect influences of each of the component characteristics of this trait. Path coefficient
analysis, which is simply a standardized partial regression coefficient partition the correlation
in to direct and indirect effect. The use of this method requires cause and effect relationship
among the variables, and the experimenter must assign direction in the casual system based up
on priori grounds of experimental evidence (Dewey and Lu, 1959).
15
Singh et al. (1985), conducted path coefficient study in pea for ten quantitative traits. They
concluded number of pods per plant, number of seeds per pod, 100-seed weight and harvest
index are the main yield components affecting yield directly. High indirect effects were
contributed by number of branches, plant height and flowering via number of pods per plant;
by pod length via 100-seed weight and by maturity via both the component traits. Protein
content had negligible effect on seed yield.
Arshadet al. (2006), in the study of character correlation and path coefficient in soybean
found days to maturity, branches, pod length, pods and 100 seed weight had positive direct
effects on grain yield. High indirect effect was also exhibited via pod length by most of the
traits hence these characters may be given more emphasis while selecting high yielding
soybean lines. Basavaraja (2002), in the study of soybean induced mutagenesis found hundred
seed weight exerted positive direct effect on grain yield.
In parameters selection for yield improvement in French bean, Babar et al. (2002), identified
positive and significant direct effect of days to flowering onseed yield while they found
negative direct effects by days to maturity andplant height.
Shivakumar (2008) conducted a research on correlation and path coefficient analysis of some
quantitative traits in 40 genotypes of soybean and the result revealed that biological yield and
harvest index were major characters influencing seed yield directly and indirectly. The results
indicated that biological yield is responsible for manipulation of seed yield in soybean.
Basavaraja (2002), Sultana et al. (2005) and Gaikwadet al. (2007) reported direct effect of
pod length on seed yield in soybean genotypes.
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).
16
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 (Arunchalamet al., 1984). The effectiveness, however,
depends on the genetic divergence among the lines being crossed. The greater the divergence,
the greater are the chances of developing superior yielding genotypes. Mahalanobis’s D2
analysis provides a means of grouping the genotypes based upon their genetic divergence
(Mahalanobis, 1936). Thus, the estimation of divergence among the lines is of paramount
importance.
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 amount of heterotic expression of F1’s
and broad spectrum of variability in segregating populations (Norden, 1980; Raoet 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.
Clustering using D2(genetic distance) matrix is useful for analyzing the divergence of the
population to identify genotypic variability. The D2statistics measures the forces of
differentiation at intra- and inter-cluster levels and determines the relative contribution of
each component trait to the total divergence (Sharma, 1998). Clusters separated by the largest
D2(genetic distance) show the maximum divergence, while the genotypes in the same clusters
or groups are less divergent (Singh and Chaudhary, 1977). In selecting genotypes from the
already chosen groups, other important characteristics such as disease resistance, earliness,
quality or even performance of particular characters should be also considered, and selecting
one genotype from each group and testing them by different statistical analysis could prove to
be fruitful.
Jaylal (1994) carried out genetic divergence study in forty genotypes of soybean using
MahalanobisD2statistics. The Wilk's test revealed highly significant differences (D2= 242.31)
for all the characters, and he grouped the forty genotypes into 9 and 7 clusters respectively,
17
based on physiological and yield attributes. The analysis for estimating the contribution of
characters to the divergence indicated carotene content and total chlorophyll in the case of
physiological and pods/cluster, branches/plant and seed yield/plant in the case of yield
attributes contributed maximum to the total genetic divergence.
Sarma and Roy (1994) classified 42 early maturing pigeon pea genotypes on the basis of
D2analysis. The analysis of variance revealed significant differences among the 42 genotypes
for allthe characters under study, indicating considerable variation among the genotypes. The
D2valuesranged from 11.5 to 2658.6, reflecting wide diversity among the genotypes. Based on
thesevalues, they grouped the 42 genotypes into eight clusters. Cluster means for
branches/plantpods/plant, harvest index and yield/plant were conspicuous and contributing
more to the totalgenetic divergence, which was also reflected by their high coefficient of
variation.
Information on the extent of genetic diversity amongst the breeding materials is very
important in the crosses between groups with maximum genetic divergence which would be
more responsive for improvement since they are likely to produce desirable recombination
and segregation in their progenies after hybridization (Norden, 1980; Reddy et al., 1988).
Barelliet al., (2005) used 35 landraces of common bean from Brazil to study the divergence
among them. They evaluated traits like, number of days to emergence, number of days to
flowering, height of the insertion of the first pod, longitudinal length of the pods, total number
of pods/plant, number of total seeds/plant, number of seeds/pod and seed weight. The genetic
distance measurements using generalized MahalanobisD2demonstrated greater dissimilarity
between genotypes from Mesoamerica and Andean gene pools. Cluster analysis grouped the
genotypes into nine clusters; with the most similar cultivars grouped in cluster I. cluster I to V
contained landraces from Mesoamerican origin, whereas groups from VII to IX only possess
Andean origin.
18
Principal component analysis (PCA) is one of the multivariate statistical techniques which is a
powerful tool for investigating and summarizing underlying trends in complex data structures
(Legendre and Legendre, 1998). Principal component analysis reflects the importance of the
largest contributor to the total variation at each axis for differentiation (Sharma, 1998). PCA
can be used to drive a two dimensional scatter plot of individuals, such that the geometrical
distance among individuals in the plot reflect the genetic distances among them with minimal
distortion. Aggregates of individuals in such a plot will reveal sets of genetically similar
individuals (Warburton and Crossa, 2000).
The first step in PCA is to calculate Eigen values, which define the amount of total variation
that is displayed on the PC axis. The first PC summarizes most of the variability present in the
original data relative to all remaining PCs. The second PC explains most of the variability not
summarized by the first PC and uncorrelated with the first and so on (Jollife, 1986).
19
20
3. MATERIALS AND METHODS
The experiment was conducted at two locations, namely Jimma Agricultural Research Center
(JARC) and Assosa Agricultural Research Center (AARC). Jimma Agricultural Research
Center is located at 70 40’9”N latitude and 36047’6”E longitude at elevation of 1753 m. a. s. l.
in south western part of Ethiopia, 365 km away from the capital Addis Ababa. It is
categorized under tepid to cool sub-humid (H2) sub agro-ecology zone of the country. The
average annual rainfall is 1559 mm. The maximum and minimum temperatures are 26.20C
and 11.30C, respectively. The major soil types of the area are chromic Nitosols and Cambisols
in the uplands, whereas Fluvisol is the dominant soil type in the bottom land and almost all
soil types have pH less than 5 (EIAR, 2008).
In this study 49 soybean genotypes (Table 1) were obtained from different sets of soybean
variety trial conducted by soybean Breeding Section of Jimma Agricultural Research Center
(JARC); that were introduced from Asian Vegetable Research Center (AVRDC) and
Mozambique. In addition, released varieties of soybean and collections from Ethiopia were
also included in this study.
21
No. Genotype Origin/source Source trial Source of seed
1 SR-4-3 Awassa-IITA SB-RVT JARC
2 TGX1895-33F PAWE-IITA SB-RVT (MS) JARC
3 H4 IIAM-Mozambique SB-RVT JARC
4 H1 IIAM-Mozambique SB-RVT JARC
5 TGX-297-6E1 Awassa-IITA JM-Seed increase 2007 JARC
6 AGS-7-1 Awassa-IITA SB-RVT JARC
7 Crawford Jimma-IITA JM-Seed increase 2007 JARC
8 AGS-234 Awassa-IITA JM-Seed increase 2007 JARC
9 H14 IIAM-Mozambique SB-RVT JARC
10 HS-82-2136 PAWE-IITA SB-RVT JARC
11 Protana 2 Awassa-IITA JM-Seed increase 2007 JARC
12 Bossier-2 Awassa-IITA JM-Seed increase 2007 JARC
13 Essex-1 Awassa-IITA JM-Seed increase 2007 JARC
14 PR-149-81-EP7 PAWE-IITA SB-RVT (LS) JARC
15 IAC-6 IIAM-Mozambique SB-RVT JARC
16 Assosa local check-1 AARC-Ethiopia JM-Seed increase 2007 JARC
17 Clark-63K JARC-Ethiopia JM-seed increase 2007 JARC
18 PR-41-(339) PAWE-IITA SB-RVT (LS) JARC
19 TGX-1895-49-F Awassa-IITA JM-Seed increase 2007 JARC
20 Davis PAWE-IITA SB-RVT (MS) JARC
21 SR-4-1 Awassa-IITA JM-Seed increase 2007 JARC
22 IAC-11 IIAM-Mozambique SB-RVT (MS) JARC
22
Table 1 continued
23
3.3 Experimental Design, Management and Season
The trials were established in the field on the main cropping season of 2010. The experiments
were laid out in 7 X 7 simple lattice design with two replications. The plot size was three rows
of 4m length with 0.6m row spacing i.e. 4m x 3 x 0.6m = 4.8m2 . Planting was done by hand
on June 10 at Jimma and June 13 at Assosa. Seed rate was 25kg/ha. Rhizobia bacteria were
incorporated in to the soil based on the standard recommendation per hectare basis to increase
Nitrogen fixing process by the genotypes to be studied. All experimental factors were applied
uniformly to the entire plot.
The data recording for each trait were carried out as follows. Five sample plants were used for
all the characters under study.
1. Days to 50% flowering: Number of days from emergence to the day on which 50 per
cent of the plants on a plot produced flowers.
2. Days to 50% pod setting: Number of days from emergence to the day on which 50
per cent of the plants on a plot set pods.
3. Days to maturity: Number of days from sowing to the stage when 95 % of the plants
in a plot have changed the color of their pods from green to yellow.
4. Hundred seed weight (g): Weight in grams of 100 seeds at harvesting.
5. Grain yield per hectare (kg): plot yield converted to hectare yield by using the
formula Grain yield (kg/ha)= (plot yield (kg) x 10,000)/plot size in square meters.
1. Biological yield per plant (g): Recorded by weighing the total above ground yield
harvested from the sample plants at the time of harvest.
24
2. Harvest Index: To estimate the harvest index, average seed yield was divided by the
average biological yield.
3. Root to biomass ratio: Average root dry weight was divided by the average above
ground biomass dry weight and expressed in percentage.
4. Root volume: Was recorded for sampled plants using volume displacement technique.
5. Pods per plant: Total number of pods for sampled plants were counted and recorded.
6. Root dry weight (g): Was estimated after drying the roots of the sampled plants in an
oven for 24 hours at 70 degree Celsius till constant weight is achieved.
7. Pod length (cm):Exterior distance of fully matured pod from the pod apex to the
peduncle was measured in centimeters from five sample plants.
8. Plant height (cm): The height of the plant from the ground surface to the tip of the
main guide was recorded in centimeters at harvesting period.
9. Total nodules per plant: The total number nodules produced for sampled plants were
counted and recorded.
10. Effective nodules per plant: Nodules that have pink color which is an indication of
fixed atmospheric nitrogen to the plant roots were counted and recorded.
The data were subjected to statistical analysis of variance as per the simple lattice design for
each character by the GLM and ANOVA procedures of SAS (SAS, version 9.2).
Efficiency of the simple lattice design relative to RCBD was checked and in most of the response
variables the lattice was found to be more efficient than that of the RCBD. In addition, test of
homogeneity of error variance was done using F-test and the result demonstrated heterogeneous error
25
variance for most of the characters; therefore, the results of the two locations were interpreted and
discussed separately. LSD was used to separate the means. Skeleton of ANOVA for simple lattice is
given below (Table 2).
Total ( r ) ( k2 ) -1
In order to identify and ascertain the genetic variability among genotypes, for the characters
under study and to confirm the presence of environmental effect on various characters,
different genetic parameters were estimated by adopting the following formulas.
Genotypic and phenotypic variances and coefficients of variation were estimated based on the
formula suggested by Burton and Devane (1953).
Msg − Mse
Genotypic variance δ g=
2
Where, r = number replication, MSg = mean square due to genotypes MSe = mean square of
error (Environmental variance)
26
Phenotypic variation δ 2 P = δ 2 g + Mse
Where, σ2g = genotypic variance, σ2p= phenotypic variance
δ 2g
Genotypic coefficient of variation GCV = ∗100
Χ
δ 2P
Phenotypic coefficient of variation PCV = ∗ 100
X
−
Where, Χ the grand mean of a character.
Broad-sense heritability (h2) for all traits was calculated using the method suggested by
Falconer (1989).
σ 2g
H = × 100
σ 2P
= genotypic variance
= phenotypic variance
The method described by Johnson et al. (1955) was followed to compute expected genetic
advance (GA).
27
GA = K *σp *H
Genetic advance as percent of mean was computed to compare the extent of predicted
advance of different traits under selection using the following formula:
GA
GAM = *100
Χ
Phenotypic correlation, the observable correlation between variables, which is the sum of
genotypic and environmental effects was calculated from variance covariance components
using the formula of Miller et al. (1958) as follows
28
Genotypic correlationis also given by
The significances of the correlation coefficients were tested using ‘r’ tabulated value at n-2
degrees of freedom, at 5% and 1% probability levels, where, n= number of treatments
(genotypes).
Based on genotypic correlation, path coefficient which refers to the direct and indirect effects
of the yield attributing traits (independent character) on grain yield (dependent character) was
estimated with the method described by Dewey and Lu (1959) as follows:
Where, rij = mutual association between the independent character (i) and dependent
character (j) as measured by the genotypic correlation coefficient.
Pij = direct effects of the independent character (i) on the dependent character (j) as measured
∑rikpkj = Summation of components of indirect
by the genotypic path coefficients, and
effects of a given independent character (i) on a given dependent character (j) via all other
independent characters (k).
29
is the direct effect of yield by ithtrait, and
Clustering of genotypes in to different groups was carried out by average linkage method. The
appropriate number of clusters was determined from the values of Pseudo F and Pseudo
T2 statistics using the procedures of SAS computer software version 9.2 facilities so as to
group sets of genotypes in to homogenous clusters (SAS 2008).
-1
D2ij = (Xi- Xj) S (Xi – Xj)
th th
Xi and Xj = the difference in mean vectors of i and j genotypes, and
-1
S = the inverse of pooled variance covariance matrix.
Principal component analysis (PCA) was used to find out the characters, which accounted
more to the total variation (Wiley, 1981). The data were standardized to mean of zero and
variance of one before computing principal component analysis. Principal component analysis
was performed using correlation matrix by employing the procedure prin comp corr of SAS
30
software version 9.2 (SAS Institute, 2001) in order to examine the relationships among 15
quantitative traits that are correlated among each other by converting in to uncorrelated traits
called principal components (PC).
31
4. RESULTS AND DISCUSSION
Results on variability assessment, associations among yield and yield related characters and
genetic divergence are presented and discussed hereunder.
Range and mean values of the 15 characters are shown in Tables 5 and 6 for Jimma and
Assosa, respectively. The mean performance of the 49 genotypes for 15 traits is presented in
Appendix Tables III and IV for Jimma and Assosa, respectively. The 49 soybean genotypes
showed wide range of variability for all characters; except pod length and root dry weight at
both locations (Table 5 and 6).
32
At Jimma, the highest grain yield (3868 kg/ha) was recorded from TGX-297-6E-1 followed
by G01892 (3735kg/ha) which were higher than the grand mean of the genotypes studied
(2160.00 kg/ha). While low yield (795 kg/ha) was obtained from the genotype PR-160-6.
At Jimma, about 57.14 per cent of the genotypes gave above the grand mean of grain yield.
If the breeding objective is to improve seed yield, genotypes with high yield in this study need
further work.
At Assosa, the highest grain yield (2134 kg/ha) was recorded from the genotype TGX-1895-
33F and the lowest yield (444 kg/ha) was obtained from G00386. The grand mean of grain
yield at Assosa was 885.76 kg/ha. Earlier days to flowering, days to pod setting and days to
maturity was observed at Assosa as compared to Jimma with mean values of (37.00), (65.50)
and (94.50) days for the genotypes F81-7636-4, FB1-7636 and H2, respectively. 46.93 per
cent of the genotypes gave above the grand mean of grain yield at Assosa.
The characters such as plant height, pod number per plant, total nodules per plant, effective
nodules per plant, biomass yield, root dry weight and harvest index showed relatively higher
values at Jimma as compared to those recorded at Assosa with mean of 76.79cm, 46.44,
31.53, 18.29, 32.99g, 8.02g, and 38.12, respectively. The genotype G01853 had scored the
highest plant height (118.17cm) at Jimma. Therefore, when breeding for higher plant height
this genotype should be considered. PR-160-6 scored the highest pod number per plant
(79.00) at Jimma indicating it could be the preferable genotype in breeding for high number
of pods per plant.
Characters such as root to biomass ratio, root volume, pod length and hundred seed weight
showed relatively higher values at Assosa compared to those at Jimma with mean values of
25.50%, 4.96, 4.49cm and 13.68g, respectively. The genotypes such as H18, Promoveria and
IAC-6 scored the highest values of root to biomass ratio (32.29%), root volume (12.50cm3)
and hundred seed weight (21.50g), respectively at Assosa. Therefore, if the breeding objective
is to improve the above traits the respective genotypes should be given due attention.
33
Table 3. Analysis of variance for 15 characters in 49 soybean genotypes tested at Jimma
(2010/2011)
Mean squares
Degrees 1 48 12 36 48
of
freedom
34
Table 4. Analysis of variance for mean square of the 15 characters of 49 soybean genotypes
tested at Assosa (2010/2011)
Mean squares
35
Table 5. Range, mean, variance, broad sense heritability, genotypic and phenotypic coefficient of variation and genetic advance as per
cent of mean for characters of soybean genotypes studied at Jimma (2010/11)
Characters Range Mean ± S.E Mean σ 2g σ 2e σ 2p GCV (%) PCV (%) H2 (%) GA GA (%)
DF 46.00-86.00 64.20±2.55 43.79 7.26 51.05 10.30 11.12 85.77 12.82 19.97
DPS 57.00-97.00 75.62±2.39 44.95 5.66 50.61 8.86 9.40 88.81 13.47 17.81
DM 111.00-148.00 125.74±2.69 42.89 8.04 50.93 5.20 5.67 84.21 13.08 10.40
PlH 37.00-120.00 76.78±4.23 240.51 14.97 255.48 20.20 20.81 94.14 32.04 41.73
PPP 26.00-79.00 46.43±4.00 137.30 14.95 152.25 25.23 26.57 90.18 23.70 51.02
PL 3.00-5.00 4.10±0.22 0.24 0.05 0.29 11.83 13.03 82.45 0.93 22.64
BY 19.00-51.00 32.99±3.22 62.48 11.07 73.55 23.96 26.00 84.94 16.20 49.11
TNPP 13.00-98.00 31.53±1.88 175.92 3.42 179.34 42.07 42.47 98.09 27.72 87.91
ENPP 7.00-66.00 21.18±1.91 104.64 3.58 108.22 48.28 49.10 96.69 21.20 100.06
RDW 2.00-5.00 2.92±0.66 0.11 0.48 0.59 11.35 26.30 18.64 0.31 10.67
RV 4.00-16.00 8.02±2.01 0.37 4.85 5.22 7.58 28.48 7.08 0.87 10.81
RBR 4.34-16.00 8.93±1.96 2.67 3.83 6.50 18.27 28.53 41.03 2.99 33.51
HSW 8.00-22.00 12.35±1.97 2.72 3.93 6.65 13.35 20.88 40.86 2.28 18.46
HI 16.80-72.63 38.21±6.38 101.82 43.31 145.14 26.41 31.53 70.15 18.65 48.80
GY 795.00-3868.00 2160.12±14.09 399246.81 189.18 399436.00 29.25 29.26 84.45 1428.4 66.14
DF=Days to 50% flowering, DPS=Days to 50% pod setting, DM=Days to maturity, PlH=Plant height, PPP=Pod per plant, PL=Pod length,
BY=Biomass yield, TNPP=Total nodules per plant, ENPP=Effective nodules per plant, RV=Root volume, RDW= Root dry weight,
RBR=Root to biomass ratio, HSW=Hundred seed weight, HI=Harvest index, GY=Grain yield. S.E. Mean= Standard error of the mean, σ
2
g= Genotypic variance, σ 2e = Environmental variance, σ 2p= Phenotypic variance, H2 (%) = Broad sense heritability, GCV (%) =
Genotypic coefficient of variation, PCV (%) = Phenotypic coefficient of variation, (%) ECV= Environmental coefficient of variation, GA=
Genetic advance, GA (%) = Genetic advance as per cent of mean.
36
Table 6. Range, mean, variance, broad sense heritability, genotypic and phenotypic coefficient of variation and genetic advance as per
cent of mean for characters of soybean genotypes studied at Assosa 2010/11
Characters Range Mean ± S.E Mean σ 2g σ 2e σ 2p GCV (%) PCV (%) H2 (%) GA GA (%)
DF 36.00-75.00 50.06±0.95 245.38 0.95 246.33 31.29 31.35 89.99 23.62 47.18
DPS 65.00-103.00 72.60±0.80 82.51 0.58 83.09 12.51 12.56 99.30 13.63 18.77
DM 94.00-120.00 111.10±0.87 135.22 0.76 135.98 10.47 10.50 99.44 16.97 15.28
PlH 20.00-62.00 35.27±1.18 197.7 1.48 199.18 39.87 40.02 99.26 20.62 58.48
PPP 9.00-28.00 15.98±0.91 35.06 0.84 35.90 37.05 37.49 97.66 8.52 53.29
PL 3.50-5.00 4.49±0.26 0.43 0.07 0.50 14.52 15.67 85.86 0.82 18.24
BY 11.00-31.00 14.12±1.33 16.49 1.82 18.31 28.96 30.52 90.06 3.11 22.20
TNPP 4.00-47.00 15.37±0.99 140.08 1.04 141.12 77.02 77.30 99.26 17.92 116.64
ENPP 2.00-25.00 7.79±1.07 49.11 1.25 50.36 90.00 91.14 97.52 10.42 133.86
RV 3.00-13.00 4.96±0.58 4.23 0.36 4.59 41.47 43.20 92.16 2.96 59.76
RDW 3.00-5.00 3.53±0.47 0.38 0.2 0.58 17.87 22.08 65.52 0.29 8.28
RBR 11.11-33.33 25.50±1.93 23.63 3.67 27.30 19.06 20.49 86.55 5.86 22.98
HSW 8.00-22.00 13.68±1.05 13.20 1.17 14.37 26.55 27.70 91.86 5.13 37.48
HI 16.13-71.07 32.79±1.37 262.38 2.08 264.46 49.39 49.59 81.25 23.86 72.74
GY 444.00-2134.00 885.76±30.51 152972.8 979.09 153951.89 44.16 44.30 82.36 593.49 67.01
DF=Days to 50% flowering, DPS=Days to 50% pod setting, DM=Days to maturity, PlH=Plant height, PPP=Pod per plant, PL=Pod length,
BY=Biomass yield, TNPP=Total nodules per plant, ENPP=Effective nodules per plant, RV=Root volume, RDW= Root dry weight,
RBR=Root to biomass ratio, HSW=Hundred seed weight, HI=Harvest index, GY=Grain yield. S.E. Mean= Standard error of the mean, σ
g= Genotypic variance, σ 2e = Environmental variance, σ 2p= Phenotypic variance, H2 (%) = Broad sense heritability, GCV (%) =
2
Genotypic coefficient of variation, PCV (%) = Phenotypic coefficient of variation, (%) ECV= Environmental coefficient of variation, GA=
Genetic advance, GA (%) = Genetic advance as per cent of mean.
37
4.2.2. Estimates of genetic parameters
Estimates of phenotypic (σ2p), genotypic (σ2g) and environmental (σ2e) variances and
phenotypic (PCV) and genotypic coefficients of variation (GCV) are provided in (Tables 5
and 6) for Jimma and Assosa, respectively.
At Jimma, grain yield, biomass yield, number of pods per plant, plant height, total nodules per
plant, effective nodules per plant, days to 50% flowering, days to pod setting, days to maturity
and harvest index exhibited high genotypic and phenotypic variances. Phenotypic coefficients
of variation (PCV) values ranged from 5.67% for days to maturity to 49.10% for effective
nodules per plant, whereas the genotypic coefficients of variation (GCV) ranged from 5.20%
for days to maturity to 48.28% for effective nodules per plant. In addition, PCV values were
generally higher than their corresponding GCV values for all the characters considered
indicating the importance of environment in the expression of these traits.
According to Deshmukhet al. (1986), PCV and GCV values greater than 20% are regarded as
high, whereas values less than 10% are considered to be low and values between 10% and
20% to be medium. Based on this delineation, PCV and GCV values were high for grain
yield, biomass yield, number of pods per plant, plant height, total nodules per plant, effective
nodules per plant, and harvest index. It indicates that selection may be effective based on
these characters and there phenotypic expression would be a good indication of genetic
potential. This result is in harmony with the findings of Jagdishet al. (2000), Patel et al.
(1998), Singh et al. (2000), Dixit (2002), and Jains and Ramgiry (2000), where high
genotypic and phenotypic coefficients of variation for grain yield, biomass yield, number of
pods per plant, plant height, and harvest index in soybean genotypes were reported. The PCV
and GCV values were medium for pod length, and days to 50% flowering. The low PCV and
GCV values were obtained for days to pod setting and days to maturity indicating lack of
adequate variability for these traits which hinders the breeding work for the improvement of
38
these traits. A low genotypic and phenotypic coefficient of variation for days to maturity is in
agreement with the reports of Ramanaet al. (2000) and Agarwalet al. (2001).
At Assosa, grain yield, days to 50% flowering, pod number per plant, plant height, total
nodules per plant, effective nodule per plant, harvest index, biomass yield, root volume and
hundred seed weight have exhibited high genotypic and phenotypic variances. Phenotypic
coefficient of variation (PCV) values ranged from 10.50% for days to maturity to 91.14% for
effective nodules per plant; whereas the genotypic coefficient of variation (GCV) ranged from
10.47% for days to maturity to 90.00% for effective nodules per plant. High PCV and GCV
values were recorded for grain yield, days to flowering, biomass yield, number of pods per
plant, hundred seed weight, plant height, root volume, total nodules per plant, effective
nodules per plant, and harvest index indicating the availability of adequate variability for
these traits which aids in the improvement of the respective characters.
39
4.2.2.2. Estimation of broad-sense heritability and genetic advance
Heritability estimate for characters under study at Jimma and Assosa are indicated in Table 5
and 6, respectively.
At Jimma, estimates of heritability in broad sense ranged from 7.08% for root volume to
99.95% for grain yield (Table 5). Similarly, root dry weight and grain yield had moderate and
high heritability with 65.52% and 99.36% respectively at Assosa. Ojo (2003), Jain and
Ramgiry (2000), and Basavaraja (2002) reported high heritability for grain yield in soybean
genotypes. According to Singh (2001), heritability values greater than 80% are very high,
values from 60-79% are moderately high, values from 40-59% are medium and values less
than 40% are low. Accordingly at Jimma, all the characters except root dry weight, root
volume, root to biomass ratio and hundred seed weight had high to very high heritability.
At Assosa, all characters except root dry weight (65%), had very high heritability. This
indicates that selection will be the best approach to be employed to identify the best soybean
genotypes for the traits with high heritability. This is because; there will be a close
correspondence between the genotype and the phenotype of the genotypes, due to the relative
small contribution of the environment to the phenotype. But, for characters with low
heritability, say 40% or less, selection may be considerably difficult or virtually impractical,
due to the masking effect of the environment. In contrary to the present study Basavaraja
(2002) reported low heritability for pod length in soybean genotypes. Low heritability values
were recorded for root dry weight and root volume at Jimma. Root dry weight showed
moderate heritability at Assosa. The magnitudes of heritability for most of the quantitative
characters at both locations were moderate to high, except the low heritability of root dry
weight and root volume at Jimma.
At Jimma, genetic advance as percent of mean ranged from 10.48 for days to maturity to
100.06 for effective nodules per plant (Table 5). At this location relatively high genetic
40
advance as percent of mean was recorded for effective nodules per plant (100.06%), total
nodules per plant (87.91%), grain yield (66.14%), pod number per plant (51.02%), biomass
yield (49.11%), harvest index (48.80%), plant height (41.73%), root to biomass ratio
(33.51%) and pod length (22.64%). Medium genetic advance as percent of mean was
recorded for days to 50% flowering (19.97%), hundred seed weight (18.46), days to pod
setting (17.81%), root volume (10.81%), root dry weight (10.67%) and days to maturity
(10.40%).
At Assosa, genetic advance as percent of mean ranged from 8.28 for root dry weight to 133.86
for effective nodules per plant (Table 6). Within this range, a relatively high genetic advance
was observed for effective nodules per plant (133.86%), total number of nodules per plant
(116.64%), harvest index (72.74%), grain yield (67.01%), root volume (59.76%), plant height
(58.48) and days to 50% flowering (47.58). Low genetic advance as per cent of mean values
were observed for root dry weight (8.28), days to maturity (15.28), pod length (18.24%) and
days to pod setting (18.77%). This low estimate of genetic advance as a percent of mean
arises from low estimate of phenotypic variance and heritability.
According to Johnson et al. (1955), high heritability estimates along with the high genetic
advance is usually more helpful in predicting gain under selection than heritability estimates
alone. The present study showed high heritability coupled with high expected genetic advance
as per cent of mean for effective nodules per plant, total nodules per plant, pod number per
plant and harvest index across both locations. These characters were controlled by additive
gene effects and phenotypic selection would likely be effective than other characters
measured (Sumati and Muralidharan, 2009).
41
4.3. Association Studies
At Jimma (Table 7) grain yield showed negative and highly significant association both at
phenotypic and genotypic levels with days to 50% flowering, days to pod setting and days to
maturity. In support of the present finding, Adityaet al., (2011), Koleet al., (2008), Ramtekeet
al. (2010) and Arshadet al., (2006), reported negative correlation of grain yield with days to
50% flowering and days to maturity in soybean genotypes. This demonstrates that whenever
the value of these characters increases, it adversely affects grain yield. This may be related to
the fact that when days to maturity increases, the phenology of the crop enters into the dry
spell, which in turn leads to decrease in yield. In contrast to the present finding Mukhekaret
al. (2004), Basavaraja (2002) and Ojwang (2003) reported positive significant association of
days to flowering with grain yield in soybean genotypes. Grain yield displayed positive non-
significant correlation with pod length, total nodules per plant, effective nodules per plant,
root dry weight, root to biomass ratio, hundred seed weight and harvest index at genotypic
and phenotypic levels. However, yield had negative non-significant association with plant
height pod number per plant and biomass yield at genotypic and phenotypic levels. Moreover,
grain yield had negative non-significant association with root volume at genotypic level.
At Assosa, grain yield manifested positive and highly significant association with pod number
per plant, biomass yield and hundred seed weight at phenotypic and genotypic level (Table 8).
Moreover, grain yield had positive significant association with root dry weight at genotypic
level. Therefore, improving one or more of the characters could result in high grain yield in
the soybean genotypes. This result is in harmony with the report of Qi Yang and Jinling Wang
(2000), Parameshwar (2006), Rajannaet al. (2000) and Malik et al., (2006).
Similar to the present study Ojo (2003) and Shivakumar (2008), reported significantly
positive correlation of grain yield with pod number per plant in soybean genotypes studied at
phenotypic level. Furthermore, grain yield had positive non-significant correlation with days
to 50% flowering, days to pod setting, plant height, root volume, harvest index and total
42
nodules per plant both at genotypic and phenotypic levels. Root dry weight had positive non-
significant correlation with grain yield at phenotypic level. Grain yield had negative non-
significant correlation with pod length and effective nodules per plant at phenotypic and
genotypic levels. This suggests the improvement of characters which had non-significant
association with grain yield will not have a sound effect on the improvement of grain yield in
the soybean genotypes. Supportive to the previous findings of Kalaimagal (1991), grain yield
had negative and significant association with days to maturity both at genotypic and
phenotypic level. The negative correlation of grain yield with days to maturity and root to
biomass ratio implies the improvement of one character affects the others in the opposite
direction making it impractical to improve the characters simultaneously. In contrary to the
present study Raman et al. (2000) and Bangaret al. (2003) reported positive significant
correlation of grain yield with days to maturity.
43
Table 7.Genotypic (above diagonal) and phenotypic correlation coefficients at Jimma (2010/11)
DF=Days to 50% flowering, DPS=Days to 50% pod setting, DM=Days to maturity, PlH=Plant height, PPP=Pod per plant, PL=Pod
length, BY=Biomass yield, TNPP=Total nodules per plant, ENPP=Effective nodules per plant, RV=Root volume, RDW= Root dry
weight, RBR=Root to biomass ratio, HSW=Hundred seed weight, HI=Harvest index, GY=Grain yield.
44
Table 8.Genotypic (above diagonal) and phenotypic correlation coefficients at Assosa (2010/11)
DF=Days to 50% flowering, DPS=Days to 50% pod setting, DM=Days to maturity, PlH=Plant height, PPP=Pod per plant, PL=Pod
length, BY=Biomass yield, TNPP=Total nodules per plant, ENPP=Effective nodules per plant, RV=Root volume, RDW= Root dry
weight, RBR=Root to biomass ratio, HSW=Hundred seed weight, HI=Harvest index, GY=Grain yield.
45
4.3.1.1. Phenotypic correlation
At Jimma, biomass yield had positive and significant phenotypic correlation with plant height
and pod number per plant. However, it had negative significant phenotypic association with
harvest index and root to biomass ratio. The positive and significant association of biomass
yield with plant height and pod number per plant as indicated in the phenotypic correlation
and among each other, indicated that these traits can be improved simultaneously through
selection. Days to 50% flowering showed positive and highly significant phenotypic
correlation with days to pod setting, days to maturity, and plant height. However it had
negative and highly significant phenotypic correlation with total nodules per plant and
effective nodules per plant. Ramanaet al. (2000), Devvartet al., (2005) and Manasa (2008)
reported significant positive association of days to 50% flowering with days to maturity in
soybean.
Days to pod setting had positive and highly significant phenotypic correlation with days to
maturity and plant height. However, it had negative and significant association with total
nodules per plant and effective nodules per plant.
Effective nodules per plant had positive and significant phenotypic correlation with total
nodules per plant. Pod number per plant showed significantly positive and negative
phenotypic association with harvest index and root to biomass ratio, respectively. In contrary
to the present finding Mukhekar (2004), Ramanaet al. (2000) and Devvartet al., (2005)
reported positive significant association of pod number per plant with harvest index.
At Assosa, days to maturity showed positive and significant phenotypic correlation with plant
height, days to 50% flowering and days to pod setting. However, it had negative and
significant association with pod per plant and harvest index. This result is in harmony with the
finding of DevVartet al., (2005) where negative association of days to maturity with harvest
index is reported. In contrary to the present study Mukhekar (2004) reported negative
significant association of days to maturity with plant height in soybean genotypes.
46
Total nodule per plant correlated positively and significantly with effective nodules per plant,
plant height, days to 50% flowering and biomass yield, it had negative and significant
association with root to biomass ratio. Days to pod setting had positive and significant
association with plant height, days to 50% flowering and days to maturity.
Biomass yield showed positive and significant correlation with total nodules per plant,
effective nodules per plant, root volume, root dry weight, hundred seed weight and plant
height. While it showed negative and significant association with root to biomass ratio.
Kausar (2005) from the study on genetic variability of F3 populations of two crosses
involving three diverse parents of soybean reported positive and significant phenotypic
correlation of biomass yield with plant height.
Plant height had positive and significant association with effective nodules per plant, days to
50% flowering, biomass yield, while it had negative significant association with root to
biomass ratio. This finding is in harmony with the report of Manasa (2008), where positive
and significant association of plant height with days to flowering is reported.
Pod number per plant had positive and negative significant association with harvest index and
days to maturity, respectively. In contrary to the present study Ramgiry and Raha (1999)
reported significantly negative association of number of pods per plant with harvest index at
phenotypic level in 64 soybean genotypes studied. Root dry weight had positive and
significant association with root to biomass ratio.
At Jimma, biomass yield showed positive and significant association with root volume, plant
height, pod number per plant, root dry weight and hundred seed weight; whereas, it had
negative and significant correlation with harvest index and root to biomass ratio.
Days to 50% flowering showed positive and highly significant correlation with days to pod
setting, days to maturity, plant height and pod number per plant. Days to 50% flowering
showed negative and significant correlation with total nodules per plant and effective nodules
per plant.
47
Days to pod setting had positive and highly significant correlation with days to maturity, root
dry weight and plant height. However, it had negative and significant association with total
nodules per plant and effective nodules per plant. Days to maturity had significantly positive
correlation with plant height. However, it had negative and significant association with pod
number per plant, root dry weight, effective nodules per plant and harvest index.
Plant height had significant and negative correlation with pod length, total nodule per plant
and root to biomass ratio. Supportive to the present study Manasa (2008), reported negative
and significant association of plant height with pod length in ovate leaflet type of soybean
genotypes. Plant height had positive significant correlation with biomass yield.
Effective nodules per plant had positive and significant correlation with total nodules per
plant. Pod number per plant had positive and significant correlation with root volume,
biomass yield and harvest index. It also showed negatively significant genotypic correlation
with root to biomass ratio.
At Assosa, days to maturity showed positive and significant correlation with plant height;
while it showed negative and significant correlation with pod number per plant, root dry
weight and harvest index. This finding is in contrary to the report of Manasa (2008) where
days to maturity had positive and significant association with pod number per plant and
harvest index.
Total nodules per plant showed positive and significant correlation with effective nodules per
plant, days to 50% flowering and plant height. However, it had negative significant
association with root to biomass ratio. Biomass yield had positive and significant association
with total nodules per plant, effective nodules per plant, root volume, hundred seed weight,
plant height, root dry weight and days to 50% flowering; while it showed negative and
significant association with root to biomass ratio and harvest index.
Plant height had positive and significant correlation with days to pod setting, biomass yield,
effective nodules per plant and root volume. However, it had negative significant correlation
48
with root to biomass ratio. Similar to the present study Gadde (2006), reported positive
significant genotypic correlation of plant height with days to pod setting in the same crop.
Effective nodules per plant showed positive and significant correlation with biomass yield and
total nodules per plant while it showed negative and significant correlation with root to
biomass ratio. Pod number per plant showed positive and significant correlation with harvest
index. Pod number per plant showed negative significant association with days to 50%
flowering and days to maturity. Root to biomass ratio had negative and significant correlation
with plant height, biomass yield and hundred seed weight. Root dry weight had positive and
significant correlation with biomass yield, root volume and root to biomass ratio.
Correlation analysis describes merely the mutual relationship between different pairs of
characters without providing the nature of the cause and effect relationships of each character.
Hence, the phenotypic and genotypic correlations were further analyzed by path coefficient
analysis technique to partition the correlation coefficient into direct and indirect effects. This
allows separation of the direct influence of each component on grain yield production from
the indirect influences caused by the mutual relationships among them. Such analysis leads to
identification of important traits useful for indirect selection of complex trait such as grain
yield (Dewey and Lu, 1959).
The Genotypic direct and indirect effect of different characters on seed yield is presented in
Tables 9 and 10 for Jimma and Assosa, respectively. At Jimma, effective nodules per plant
had the highest positive direct effect (0.837). Moreover, the indirect effect via other traits was
negative and hence the correlation it had with yield was largely due to the direct effect. This
49
suggests the correlation showed the true relationship and direct selection through this
character will be effective (Singh and Chaudhary, 1979).
The high positive direct effect of hundred seed weight on grain yield was counter balanced by
indirect negative effects of root to biomass ratio, total nodules per plant, root dry weight and
harvest index and reduced the correlation to 0.120.
Total nodules per plant which had the highest negative direct effect revealed prominent
indirect effects via days to maturity, effective nodules per plant, days to 50% flowering, days
to pod setting, pod length, biomass yield and root volume.
The third highest positive direct effect of root to biomass ratio was counter balanced by
indirect negative effects of total nodules per plant, pod number per plant, harvest index, plant
height and root dry weight and reduced the correlation to 0.190.
Direct positive effect of pod length on grain yield was counter balanced by indirect negative
effects of days to maturity, plant height, root to biomass ratio, biomass yield, total nodules per
plant, and hundred seed weight and reduced the correlations to 0.08.
The second highest negative direct effect of harvest index (-0.459) is counter balanced by the
indirect positive effect via pod number per plant, biomass yield, root volume, root to biomass
ratio, effective nodules per plant, days to maturity and days to pod setting and reduced the
correlation to 0.120.
The indirect negative effect of pod number per plant via days to 50% flowering, days to pod
setting, days to maturity, harvest index, biomass yield, pod length, root to biomass ratio and
harvest index counter balanced the positive direct effect of pod number per plant on grain
yield and reduced the correlations to –0.051. The correlation of pod number per plant with
grain yield was negative and path analysis showed that the negative correlation was mainly
due to the indirect negative effect of the character.
50
Even though root volume had direct negative effect (-0.269) on grain yield, it was counter
balanced by the indirect positive effect via the characters root dry weight, harvest index, total
nodules per plant, pod length and plant height as a result the correlation was changed to 0.011.
Path analysis showed the positive correlation of root volume with grain yield was through
indirect effects of other characters.
The direct positive effect of plant height (0.290) was counter balanced by indirect negative
effects of days to maturity, root to biomass ratio, biomass yield, days to pod setting, effective
nodules per plant, days to 50% flowering, pod length, root volume and root dry weight. The
negative correlation of this character was mainly due to the indirect negative effect of other
characters.
The positive direct effect of root dry weight (0.214) was counter balanced by the negative
indirect effect of root volume, hundred seed weight, root to biomass ratio, effective nodules
per plant, biomass yield, plant height, days to maturity, days to flowering and days to pod
setting and the correlation was changed to -0.520. Path analysis showed that the negative
correlation of root dry weight with grain yield was due to the indirect effects of other
characters.
The second highest negative direct effect of days to maturity (-0.615) was counter balanced
by positive indirect effect of total nodules per plant, plant height, pod number per plant, root
dry weight, hundred seed weight and harvest index, which reduced the correlation to -0.52.
Path analysis showed that increase in days to maturity will negatively affect grain yield.
Days to 50% flowering, days to pod setting, days to maturity and biomass yield had negative
direct effect on grain yield indicating any increase in these characters affects grain yield in the
negative direction. Therefore, selecting genotypes having less number of days to flowering,
less number of days to pod setting and less number of days to maturity could be used to
improve seed yield in soybean genotypes, as a result of their direct effect on yield.
51
The residual effect (0.252) indicated that characters which are included in the genotypic path
analysis explained (74.80%) of the total variation in grain yield which indicates that there may
be some more components that are contributing towards seed yield.
Path analysis at Jimma indicated selecting genotypes having high root to biomass ratio,
hundred seed weight and effective nodules per plant could be used to improve seed yield in
soybean genotypes as a result of their direct effect on grain yield.
52
Table 9. Path coefficients of direct (bold diagonal) and indirect effects (off diagonal) at genotypic level of 15 traits on grain yield on
49 soybean germplasm tested at Jimma (2010/11).
DF=Days to 50% flowering, DPS=Days to 50% pod setting, DM=Days to maturity, PlH=Plant height, PPP=Pod per plant, PL=Pod
length, BY=Biomass yield, TNPP=Total nodules per plant, ENPP=Effective nodules per plant, RV=Root volume, RDW= Root dry
weight, RBR=Root to biomass ratio, HSW=Hundred seed weight, HI=Harvest index.
53
At Assosa (Table 10) pod number per plant had the highest positive direct effect (0.792) on
grain yield followed by hundred seed weight (0.546), biomass yield (0.227), days to 50%
flowering (0.180) and plant height (0.094). Path analysis showed that the positive and
significant correlation of pod number per plant with grain yield was the true relationship.
Similar results were reported by Haghiet al., (2011), Gupta (2008) and Arshadet al, (2006) on
the same crop. In contrary to the present result Iqbalet al, (2003) and Agdew and Getnet
(2005) reported negative direct effect of pod number per plant on grain yield of soybean
genotypes.
The highest negative direct effect of harvest index (-0.536) was counter balanced by the
favorable indirect effect of pod number per plant, hundred seed weight, days to maturity and
total nodules per plant and the correlation was reduced to 0.250. This finding is in harmony
with the report of Agdew and Getnet (2005) and Haghiet al, (2011) where harvest index had
negative direct effect on soybean yield. In contrary to the present finding Shivakumar (2008)
reported positive direct effect of harvest index on grain yield of soybean.
The second highest positive direct effect of hundred seed weight (0.546) was counter
balanced by the indirect negative effects of the characters biomass yield, root to biomass ratio,
effective nodules per plant, pod number per plant, pod length, plant height, days to pod setting,
and days to 50% flowering. Similar results were reported by Agdew and Getnet (2005),
Shivakumar (2008) and Arshadet al, (2006) in soybean genotypes. Path analysis showed that
correlation explained the true relationship of these two characters. In contrary to the present
finding Haghiet al, (2011) and Gupta (2008) reported negative direct effect of hundred seed
weight on grain yield on the same crop.
The third highest positive direct effect of biomass yield (0.227) showed significant positive
correlation with grain yield and path analysis showed that the correlation explained the true
relationship of the character with grain yield. This result is in harmony with the findings of
Showcat and Tyagi (2006) and Shivakumar (2008) where positive direct effect of biomass
yield is reported in soybean genotypes.
54
However, root dry weight, root volume, total nodules per plant and days to pod setting had
showed negative direct effect on grain yield. They only contributed to grain yield mainly via
their positive indirect effect with other characters.
Days to maturity, pod length, effective nodules per plant and root to biomass ratio had
negative direct effect on grain yield. Moreover, their correlation with grain yield is negative
which suggested any increase in these characters affects grain yield in the negative direction.
Showkat and Tyagi, (2010) reported negative direct effect for days to maturity and pod length
in 40 soybean genotypes. The residual effect (0.231) indicated that characters which are
included in the genotypic path analysis explained (76.6%) of the total variation in grain yield
which indicates that there may be some more components that are contributing towards seed
yield.
Path analysis at Assosa indicated selecting genotypes having high number of pods per plant,
biomass yield and hundred seed weight could be used to improve seed yield in soybean
genotypes as a result of their direct effect on grain yield.
55
Table 10. Path coefficients of direct (bold diagonal) and indirect effects (off diagonal) at genotypic level of 15 traits on grain yield on
49 soybean germplasm tested at Assosa (2010/11).
DF=Days to 50% flowering, DPS=Days to 50% pod setting, DM=Days to maturity, PlH=Plant height, PPP=Pod per plant, PL=Pod
length, BY=Biomass yield, TNPP=Total nodules per plant, ENPP=Effective nodules per plant, RV=Root volume, RDW= Root dry
weight, RBR=Root to biomass ratio, HSW=Hundred seed weight, HI=Harvest index.Residual effect = 0.296
56
4.5. Cluster Analysis
Divergence analysis is a technique used to categorize genotypes that are similar into one
group and others into a different group. D-square statistics (D2) developed by Mahalanobis
(1936), has been used to classify the divergent genotypes into different groups. The genetic
improvement through hybridization and selection depends upon the extent of genetic diversity
between parents.
At Jimma, the genotypes were grouped in to five distinct clusters (Table 11). This indicates
the tested soybean genotypes were moderately divergent. The genotypes were distributed in
such a way that 21 (42.85%) genotypes were grouped into Cluster I, 14 (28.57%) genotypes
into Cluster III, 8 (16.32%) genotypes into Cluster II, 4 (8.16%) genotypes into cluster IV and
2 (4.08%) genotypes into cluster V.
Table 9. The distribution of genotypes into five clusters based on D2 analysis for 49 soybean
genotypes tested at Jimma (2010/11).
57
At Assosa, the genotypes were classified into three clusters (Table 12). Cluster II was the
largest cluster with 29 (59.18%)genotypes followed by cluster I which contained 19
genotypes or almost 38.77% of the total population. Cluster III contained only 1 genotype
which is 2.04% of the total population.
Table 10. The distribution of genotypes into three clusters based on D2 analysis for the 49
soybean genotypes tested at Assosa (2010/11).
III 1 TGX-1895-33F
The pair wise generalized squared distance (D2) among clusters is depicted in table 13 and 14
for Jimma and Assosa, respectively.
58
The χ2-test for the five clusters at Jimma (Table 13) indicated that there was statistically
significant difference among the clusters except between cluster I and III (22.65). The
maximum distance was found between cluster II and V (D2=305.26) followed by cluster III
and V (D2 =179.31), cluster II and cluster IV (D2 = 162.82), cluster III and IV (D2 = 90.35),
cluster I and V (D2 =87.13), cluster I and II (D2 = 77.19) and cluster IV and V (D2= 60.58).
The χ2-test for the three clusters at Assosa (Table 14) indicated that there was statistically
significant difference among the clusters except cluster I and III (11.85). The highest cluster
distance was recorded between cluster I and cluster III (D2=304.36) followed by cluster I and
cluster III (D2=224.41), which revealed that these clusters were genetically more divergent
from each other.
According to Ghaderiet al. (1984), increasing parental distance implies a great number of
contrasting alleles at the desired loci, and then to the extent that these loci recombine in the F2
and F3 generation following a cross of distantly related parents, the greater will be the
opportunities for the effective selection for yield factors.
Crosses involving parents belonging to most divergent clusters are expected to manifest
maximum genetic recombination and variation in genetic architecture (Singh et al., 1987). For
instance, in the present result (Jimma) crosses involving parents belonging to most divergent
clusters, for example clusters V with cluster II, cluster V with cluster III, and cluster IV with
cluster II are expected to provide relatively better genetic recombination and combination in
their progenies.
However, the selection of parents should also consider the special advantages of each cluster
and each genotype within a cluster depending on specific objectives of hybridization (Singh,
2001; Chahal and Gosal (2002).
59
Table 11.Mahalanobis distance between groups of soybean genotypes at Jimma
CLUSTERS I II III IV V
I - 77.19** 22.65ns 30.65** 87.13**
II - 27.17* 162.82** 305.26**
III - 90.35** 179.31**
IV - 60.58**
V -
χ2= 23.68 and 29.14 at 5%, 1% probability level respectively.
CLUSTERS I II III
I - 11.85ns 304.36**
II - 224.41**
III -
χ2= 23.68 and 29.14 at 5%, 1% probability level respectively.
Populations from areas far separated geographically and having complex environment are
normally expected to accumulate enormous genetic diversity (Chandel and Joshi, 1983).
However, the distribution of strains in different clusters did not follow definite pattern with
regard to geographical origins in the present case. Some accessions from different regions
were found to be closely related regardless of their geographic origin (source) and the rugged
nature of the terrain which could have favored isolation among the genotypes and hence,
distinct lines of evolution in each region. This could be realized from the overlapping in
clustering pattern among genotypes from different origin. In most of the cases, genotypes
from same place of origin fell in to the different clusters and from different places of origin
fell in to same cluster. Regarding to genotypes collected from Ethiopia, at Jimma, those
60
genotypes from Awassa area are distributed in to cluster I (47.36%) and the rest of the
genotypes were distributed in to cluster II to cluster V each cluster having 10.52% of the
genotypes. Genotypes from Pawe area are also distributed in to different clusters at Jimma.
For instance, 40% of the genotypes are in cluster III, 30% in cluster II, 20% in cluster I and
10% in cluster IV. The genotypes from Jimma area are distributed in cluster III and cluster IV
each having 66.66% and 33.34%, respectively. The genotype from Assosa area is found in
cluster II.
At Assosa, genotypes from Awassa area are distributed in cluster I (27.77%) and in cluster II
(72.23%). Genotypes from Pawe area are distributed in to the three clusters 60% in cluster II,
30% in cluster I and the rest 10% in cluster III. The genotypes from Jimma area are
distributed in cluster I (66.66%) and in cluster II (33.34%). The genotype from Assosa area is
found in cluster II.
Several possible reasons could be given for the genetic similarity among accessions from
different regions. There could also be a tendency, particularly among resource poor farmers in
marginal areas, of selecting for the same traits of interest like yield stability, resistance to
diseases, insects and abiotic calamities and low dependence on the external inputs (de Boefet
al,. 1996). Although the original sources might vary, the crop might have also been forced to
evolve in the same direction by this kind of local breeding for the same targets which may
emanate from similar economic, social cultural and ecological reasons in the area.
In this study the results showed that there was moderate diversity in soybean genotypes.
Genetic architecture of a population is generally believed to be the result of breeding system,
gene flow within and between populations, isolation mechanisms and prolonged selection by
various natural and artificial forces (Chandel and Joshi, 1983). Ecological environment is
believed to be the major force in crop evolution (Spagnoletti and Qualset, 1987). Therefore
this diversity in soybean genotypes could mainly be attributed to diverse agro-climatic
conditions of the areas from where they were collected (Harlan, 1969). However, there was no
definite relationship between geographic diversity and genetic diversity. It is suggested that
61
selection of parents for hybridization need not necessarily be based on geographic diversity
but genetic diversity must form the base for parental selection.
At Jimma, mean value of the 15 quantitative characters in each cluster is presented in Table
15. The characteristic feature of each cluster is discussed hereunder. The data obtained from
the two locations were checked for homogeneity following F test but they were found to be
heterogeneous therefore the results are analyzed and discussed independently.
Cluster I had medium maturity (123.9 days), high pod length (4.27cm) and heavier seed
weight (12.75g). Cluster II had late days to flowering (73.00 days), late days to pod setting
(84 days), late days to maturity (137 days), highest plant height (92.07), highest pod number
per plant (58.50), highest root dry weight (2.90g), lowest number of total nodules per plant
(17), and the lowest grain yield (1158.40 kg/ha).
Cluster III had a characteristic feature of highest biomass yield (35.00g), highest root volume
(8.93cm3), shortest plant height (72.66cm), lowest root dry weight (2.64g) and lowest root to
biomass ratio (7.97%).
Cluster IV had characteristic feature of highest number of total nodules per plant and effective
nodules per plant with 42.38 and 37.50, respectively. Moreover, the cluster also had the
highest harvest index (51.41%), highest root to biomass ratio (10.08%), the lowest biomass
yield (30.38g) and the shortest pod length of (3.69g).
Cluster V could be characterized by early flowering (55.00 days), early days to pod setting
(71.00 days), early maturity (119.65 days), lowest number of pods per plant (39.50), lowest
seed weight (11.00g), lowest harvest index (29.44%), lowest root volume (6.25cm3), lowest
number of effective nodules per plant (13.50) and highest grain yield (3795 kg/ha).
62
Table 13. Cluster mean for 15 characters in soybean tested at Jimma (2010/11)
At Assosa, cluster I had characteristics of early days to 50% flowering (48.60days), early days
to pod setting (71.52 days), highest pod length (4.56cm), lowest root dry weight (3.42cm),
lowest hundred seed weight (12.92g), lowest number of pods per plant (14.02), shortest plant
height (33.18cm) and the lowest grain yield of 630.78kg/ha (Table 16).
63
Cluster II had early days to maturity (108.87 days), the highest number of pods per plant
(17.27), the highest harvest index (36.42%), the lowest number of total nodules per plant
(14.68) , effective nodules per plant (7.24) and the lowest root volume (4.81cm3).
Cluster III is characterized by late flowering (65days), late days to pod setting (73days), and
medium maturity (116days). The cluster also had tallest plant height (61.20cm), highest
number of total nodules per plant (29.50), highest number of effective nodules per plant
(11.00), highest root dry weight (4.50g), highest biomass yield (24.00g), lowest root to
biomass ratio (15.71%), highest seed weight (19.50g), lowest harvest index (17.50%) and the
highest grain yield of (2130 kg/ha).
Table 14. Cluster mean for 14 characters in soybean tested at Assosa (2010/11)
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Grain yield 630.78* 1009.67 2130.00**
Principal component analysis (PCA) is one of the multivariate statistical techniques which is a
powerful tool for investigating and summarizing underlying trends in complex data structures
(Legendre and Legendre 1998). Principal component analysis reflects the importance of the
largest contributor to the total variation at each axis for differentiation (Sharma, 1998).
The principal component analysis at Jimma revealed that six principal components PC1 to
PC6 with Eigen values 4.20, 1.99, 1.91, 1.40, 1.29 and 1.17 respectively, have accounted for
79.90% of the total variation (Table 17).
The PC1 which accounted for 28.00% of the total variation among accessions at Jimma was
mainly due to the contrast between days to 50% flowering, days to pod setting, days to
maturity, plant height and total nodules per plant. Likewise, 13.27% of the total variation
among the tested accessions accounted for the second PC originated from the contrasting
effect between biomass yield, root volume, root to biomass ratio and harvest index.
Similarly, the third PC, which explained 12.77% of the total variation was mainly due to the
contrast between pod number per plant, effective nodules per plant, harvest index, root dry
weight and root to biomass ratio. The PC4 which explained 9.39% of the total variation
among the accessions was due to the contrast between root volume, pod number per plant and
hundred seed weight. Similarly, PC5, which accounted for 8.64% of the total variation, was
mainly due to the average effect of plant height, total nodules per plant, effective nodules per
plant and root dry weight. PC6 which explained 7.83% of the total variation among the
accessions was due to the contrast between pod length, root dry weight and grain yield.
65
At Assosa, the principal component analysis in (Table 17) revealed that five principal
components PC1 to PC5 with Eigen values 4.27, 2.53, 1.91, 1.28 and 1.08 respectively, have
accounted for 73.81% of the total variation.
The PC1 which explained 28.43% of the total variation resulted from the contrast between
biomass yield, days to 50% flowering, plant height, total nodules per plant, effective nodules
per plant and root to biomass ratio. Correspondingly, the PC2 which accounted for 16.89% of
the total variation was attributed to the contrast between days to maturity, pod number per
plant, biomass yield, hundred seed weight, harvest index and grain yield.
PC3 which explained 12.71% of the total variability among the tested accessions resulted
from the contrast between pod length, root volume, root dry weight, effective nodules per
plant and root to biomass ratio. Similarly, the PC4 which accounted for 8.54% of the total
variation among the tested genotypes was mainly due to the average effect of days to 50%
flowering and days to pod setting. The average effect of total nodules per plant, effective
nodules per plant, root dry weight and root to biomass ratio in the fifth PC, accounted for
7.23% of the total variation.
According to Chahal and Gosal (2002), characters with largest absolute values closer to unity
within the first principal component influence the clustering more than those with lower
absolute values closer to zero. Therefore, in the present study, differentiation of the genotypes
into different cluster was because of a cumulative effect of a number of characters rather to
the small contribution of each character.
66
Table 15. Eigenvectors, total variance explained, cumulative and eigen values of the first seven and six principal components (PCs) of
soybean genotypes evaluated at Jimma and Assosa respectively.
Eigen vectors
Jimma Assosa
Characters PC1 PC2 PC3 PC4 PC5 PC6 PC1 PC2 PC3 PC4 PC5
DF 0.410 -0.166 0.032 0.088 -0.061 -0.085 0.331 -0.035 0.063 0.490 0.128
DPS 0.398 -0.268 -0.022 0.085 0.024 -0.025 0.211 -0.087 0.174 0.649 -0.023
DM 0.367 -0.197 0.045 -0.140 0.209 -0.157 0.255 -0.358 0.007 0.098 -0.297
PH 0.305 -0.034 0.137 -0.296 0.389 0.162 0.344 0.096 0.081 0.152 -0.059
PPP 0.122 0.029 0.451 0.529 -0.135 0.194 -0.250 0.375 -0.060 0.190 -0.001
PL -0.085 0.254 -0.279 0.001 -0.235 -0.320 -0.107 0.060 -0.465 0.142 0.115
BY 0.254 0.556 0.148 0.022 0.077 0.160 0.329 0.309 0.117 -0.245 -0.088
TNPP -0.354 0.067 0.108 0.160 0.400 -0.256 0.372 0.055 -0.263 -0.117 0.360
ENPP -0.242 0.050 0.301 0.159 0.578 -0.128 0.338 0.002 -0.321 -0.139 0.443
RV 0.168 0.314 -0.192 0.479 -0.076 -0.124 0.183 0.137 0.380 -0.240 -0.141
RDW 0.129 0.071 -0.356 0.256 0.319 0.527 0.001 0.215 0.516 -0.017 0.474
RBR -0.199 -0.449 -0.388 0.141 0.127 0.185 -0.312 -0.165 0.302 0.090 0.471
HSW -0.029 0.149 0.267 -0.422 -0.097 0.169 0.156 0.390 -0.098 0.021 0.056
HI -0.134 -0.376 0.433 0.208 -0.273 0.094 -0.266 0.358 -0.192 0.291 0.025
GY -0.267 0.103 0.004 -0.103 -0.138 0.573 0.048 0.489 0.072 0.073 -0.274
Eigen values 4.20 1.99 1.91 1.40 1.29 1.17 4.27 2.53 1.91 1.28 1.08
Total variance explained 28.00 13.27 12.77 9.39 8.64 7.83 28.43 16.89 12.71 8.54 7.23
Cumulative 28.00 41.28 54.04 63.43 72.07 79.90 28.43 45.32 58.04 66.58 73.81
67
5. SUMMARY AND CONCLUSION
The progress of crop improvement program depends on the choice of material, the extent of
variability present and the knowledge of quantitative characters with grain yield and related traits.
The present study comprises 49 soybean genotypes that were evaluated at two locations, namely
Jimma and Assosa with the objective of assessing the genetic variability and character
associations for 15 characters.
The results of analysis of variance for each location showed the genotypes were significantly
different at (P<0.01) for all characters except root volume and root dry weight at Jimma.
Phenotypic coefficient of variability (PCV) values at Jimma ranged from 5.67% for days to
maturity to 49.10% for effective nodules per plant, whereas the genotypic coefficient of
variability (GVC) ranged from 5.20% for days to maturity to 48.28% for effective nodules per
plant. Phenotypic coefficient of variability values were low for days to pod setting, and days
to maturity; medium for pod length and days to 50% flowering and it was high for the rest of
the characters. Genotypic coefficient of variability values were low for days to pod setting,
days to maturity, and root volume; high for grain yield, biomass yield, number of pods per
plant, hundred seed weight, plant height, total nodules per plant, effective nodules per plant,
and harvest index. The high GVC values of these characters suggest the possibility of
improving these traits through selection.
At Assosa, phenotypic coefficient of variability (PCV) values ranged from 10.50% for days to
maturity to 91.14% for effective nodules per plant, whereas the genotypic coefficient of
variability (GVC) ranged from 10.47% for days to maturity to 90.00% for effective nodules
per plant. The traits such as, grain yield, days to 50% flowering, biomass yield, number of
pods per plant, hundred seed weight, plant height, root volume, total nodules per plant,
effective nodules per plant, and harvest index had high phenotypic (PCV) and genotypic
coefficient of variability (GCV) values.
High heritability coupled with high expected genetic advance (as percent of mean) was
observed for effective nodules per plant, total nodules per plant, harvest index, pod number
68
per plant and grain yield for both locations; biomass yield, plant height and root volume in
addition to the fore mentioned traits at Assosa. Thus, these characters can be improved
through selection more easily than other characters.
Correlation analysis at Jimma showed that grain yield had negative and significant association
with days to 50% flowering, days to pod setting and days to maturity both at phenotypic and
genotypic levels. At Assosa, grain yield showed positive significant association with pod
number per plant, hundred seed weight and biomass yield at genotypic and phenotypic level;
and with root dry weight at phenotypic level. Grain yield had negative and significant
correlation with days to maturity and root to biomass ratio at phenotypic level. Selecting for
those traits showing positive and significant correlation coefficient with grain yield supports
the possibility to increase grain yield of soybean.
Genotypic correlation coefficients of various characters with seed yield were partitioned into
direct and indirect effects. At Jimma, effective nodules per plant exerted the highest positive
genotypic direct effect followed by hundred seed weight, root to biomass ratio, pod number
per plant, plant height, root dry weight and pod length. The rest of the characters had negative
direct effect on grain yield. The highest direct positive effect at Assosa were exerted by pod
number per plant followed by hundred seed weight, biomass yield, days to 50% flowering,
and plant height. The rest of the characters had negative direct effect on grain yield.
Therefore, root to biomass ratio, hundred seed weight and effective nodules per plant; pod
number per plant, hundred seed weight and biomass yield at Jimma and Assosa, respectively
were the important contributors to seed yield and these traits could be used as an indirect
selection criterion.
69
Based on the relative squared distance values (D2) between any two genotypes, the 49
soybean genotypes were grouped into five and three distinct clusters at Jimma and Assosa,
respectively. This indicates that the soybean genotypes were moderately divergent.
The principal component analysis at Jimma revealed six principal components (PCs) having
eigenvalues between 1.17 and 4.20 extracted a cumulative of about 79.90% of the total
variation noted among the genotypes. It was also noted that differentiation of the genotypes
into different cluster was because of a cumulative effect of a number of characters rather than
the small contribution of each character. The principal component analysis at Assosa
indicated five principal components (PCs) having eigenvalues between 1.08 and 4.27
explained a cumulative of 73.81% of the total variation among the genotypes.
The present study generally implied the presence of significant genetic variability among the
tested genotypes. Thus, there is an opportunity to bring about improvement through direct
selection or hybridization. However, all the above conclusions were derived from results of
studies conducted within one season. So, further studies of soybean genotypes with larger
sample size in broad environments and seasons should be conducted on soybean variability in
order to give confirmative results.
70
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81
7. APPENDICES
82
Appendix Table I. Analysis of variance for mean square of the 15 characters of49 soybean
genotypes tested at Jimma (2010/2011)
Mean squares
Degrees 1 48 12 36 48
of
freedom
83
Appendix Table II. Analysis of variance for mean square of the 15 characters of49 soybean
genotypes tested at Assosa (2010/2011)
Mean squares
84
Appendix Table III. Analysis of variance (ANOVA) for the combined characters of 49
soybean genotypes tested at Jimma and Assosa (2010/2011)
Mean squares
PL= Pod length, RDW= Root dry weight, RBR= Root to biomass ratio, HSW=
Hundred seed weight
85
Appendix Table IV. Mean values of 15 traits of soybean genotypes grown at Jimma (2010/11)
86
JSL 1 83.00 97.00 135.71 71.54 48.00 4.00 35.00 22.00 16.00 4.00 9.00 11.67 10.00 28.50 795.00
G9945 67.00 75.00 122.46 66.30 45.00 5.00 21.50 17.00 4.00 2.50 8.00 11.85 11.00 51.27 2619.0
Hardee-1 60.00 70.00 125.58 74.05 38.00 4.00 30.50 28.00 14.00 3.50 8.00 11.42 12.00 32.17 2381.0
G01892 49.00 67.00 114.29 77.87 43.00 4.00 27.00 29.00 13.00 3.00 7.50 10.87 10.00 30.80 3735.0
H3 69.00 76.00 124.68 70.90 46.00 3.75 26.00 18.00 9.00 3.00 7.50 11.53 12.00 38.77 2769.0
IAC-73-5115 64.00 73.00 117.93 41.58 56.00 3.75 25.50 40.00 22.00 2.75 9.50 10.81 9.00 43.09 1610.0
PR-149-81-EP 77.00 87.00 138.05 94.66 43.00 3.75 33.50 28.00 7.00 4.00 10.50 11.96 14.00 38.91 1274.0
AGS 214 63.00 68.00 125.65 56.97 44.00 5.00 36.00 43.00 17.00 2.75 9.00 7.64 16.00 41.72 1961.0
AGS-3-1 65.00 75.00 120.40 66.73 78.00 3.75 32.00 45.50 33.00 2.50 8.00 7.81 13.00 68.19 2982.0
F81-7636-4 64.00 71.00 125.51 53.98 38.00 4.00 20.00 29.00 11.00 2.50 7.00 12.50 14.00 56.90 2658.0
G00391 48.00 59.00 116.97 66.46 29.50 4.00 21.50 94.00 60.00 2.50 6.00 11.69 14.00 41.77 2352.0
G03705 54.00 64.00 111.86 57.49 42.00 4.00 25.50 49.00 20.00 2.50 10.00 9.77 11.00 38.17 1863.0
G01853 76.00 86.00 127.12 118.17 58.00 3.00 31.00 20.50 20.00 2.50 7.00 8.02 9.00 35.63 1327.0
H10 69.00 76.00 125.23 81.25 32.00 5.00 47.00 46.00 23.00 3.00 9.00 6.39 13.00 18.66 2009.0
V1-1 65.00 71.00 124.83 71.56 37.00 4.00 43.00 36.00 15.00 2.25 7.00 5.22 21.00 41.36 2698.0
AGS-299-2 68.00 75.00 122.58 58.82 53.00 4.00 24.00 23.00 12.00 2.00 7.00 8.39 11.00 55.84 2525.0
PR-145-2 59.00 66.00 124.69 89.07 73.00 4.00 46.00 36.00 29.00 3.00 8.00 6.53 11.00 36.31 2202.0
FB1-7636 69.00 77.00 128.86 91.95 38.00 5.00 34.00 25.00 5.00 4.00 7.00 11.93 10.00 21.21 2171.0
F82-7629-2 59.00 71.00 125.75 45.97 36.00 4.00 21.00 53.00 18.00 2.50 8.00 11.82 10.00 36.68 1750.0
G00386 74.00 83.00 135.00 99.66 52.00 3.00 27.00 20.00 9.00 2.50 7.00 9.17 14.00 59.40 1571.0
AGS-3 48.00 65.50 118.12 76.73 31.00 3.75 24.00 32.00 11.00 2.50 4.50 10.31 18.00 47.89 1681.0
G00141 61.00 69.00 125.71 77.04 39.00 4.00 20.50 38.00 35.00 3.00 6.00 14.71 13.00 53.58 3100.0
H5 65.00 72.00 127.46 97.80 63.00 3.00 31.00 38.00 36.00 3.50 8.50 11.25 11.00 46.65 3146.0
Assosa local 70.00 82.00 136.81 94.41 62.00 4.75 37.00 27.00 20.00 2.50 7.00 6.70 14.00 50.84 1348.0
check-1
Mean 64.20 75.62 125.74 76.79 46.44 4.10 32.99 31.53 21.18 2.92 8.02 8.93 12.35 38.21 2160.12
CV 3.97 3.16 2.14 5.50 8.62 5.44 9.78 5.96 9.02 22.76 25.07 21.96 16.03 16.70 0.65
LSD (5%) 5.41 4.78 5.70 8.30 7.77 0.46 6.69 3.72 3.80 1.39 4.30 3.93 3.98 13.23 27.65
Appendix Table V. Mean values of 15 traits of soybean genotypes grown at Assosa (2010/11)
87
Name DF DPS DM PH PPP PL BY TNPP ENPP RV RDW RBR HSW HI GY
IAC-6 43.00 71.50 96.00 34.00 21.50 4.75 16.00 4.50 2.50 4.50 4.50 28.04 21.50 57.12 1253
SR-4-3 74.50 94.00 119.50 53.50 12.00 3.75 12.50 15.00 5.50 4.50 3.50 27.92 14.50 27.64 899.55
TGX-1895-33F 65.50 73.50 116.50 61.50 15.50 4.00 24.00 29.50 11.00 8.50 4.50 15.71 19.50 17.5 2130
H4 47.00 69.50 117.00 41.50 12.50 4.00 14.00 8.50 2.50 7.50 3.50 24.87 12.50 24.1 699
H1 42.00 71.00 114.50 21.50 16.50 4.75 13.00 9.00 3.00 4.50 3.50 26.78 11.50 28.5 581
TGX-297-6E-1 44.50 69.50 115.50 34.50 24.00 5.00 12.00 6.00 2.50 4.50 3.00 25.17 14.00 70.54 1002
AGS-7-1 69.50 89.00 115.50 23.50 13.50 5.00 14.50 5.50 2.50 4.50 3.50 24.04 10.50 22.78 964
Promoveria 41.50 72.00 116.50 35.50 18.50 4.00 13.00 4.50 3.50 12.50 4.00 30.95 13.00 37.53 569.5
Crowford 66.50 77.50 116.50 38.50 9.50 5.00 14.00 14.50 5.50 3.50 3.50 24.87 12.00 18.03 519
Lotus 41.50 69.00 98.50 32.00 27.00 4.25 13.00 28.00 11.00 4.50 3.50 26.78 12.50 55.36 1018.5
AGS-234 45.00 70.50 96.00 33.50 24.50 5.00 14.50 10.00 4.50 4.50 3.50 27.61 16.00 59.36 1169.5
H14 42.50 73.50 101.00 25.00 17.50 5.00 13.00 7.00 3.00 5.50 3.50 26.78 12.50 33.61 668.5
HS-82-2136 69.50 72.50 117.00 32.50 19.50 4.00 13.00 12.00 4.50 5.50 3.50 26.78 13.00 42.55 851
Protana 72.50 73.50 112.50 56.50 11.50 4.00 14.50 24.00 14.00 5.00 4.00 27.62 15.50 26.78 875
Essex 49.50 71.50 116.00 31.00 10.00 4.50 13.00 30.00 18.50 5.50 3.50 26.78 18.00 24.32 653
Clark-63k 41.50 69.00 115.50 22.50 12.00 4.00 13.50 7.00 3.50 4.50 3.50 29.66 12.50 21.21 780
PR-41 (339) 68.50 76.00 114.50 34.50 13.50 5.00 13.00 24.00 16.00 4.00 3.50 26.78 12.00 23.57 521
TGX-1895-49-F 66.50 73.50 116.50 52.50 16.50 4.75 14.00 27.50 15.00 4.50 3.50 24.87 14.50 33.23 917.5
Davis 42.50 67.50 95.00 26.50 18.50 3.75 12.00 5.50 2.50 3.50 3.50 29.02 12.50 37.36 945
H2 41.50 68.00 94.50 39.00 19.50 5.00 13.00 6.50 3.50 4.00 3.50 26.78 15.00 43.92 1295.5
SR-4-1 69.50 78.50 115.00 45.50 13.50 3.75 17.50 12.50 4.50 8.50 3.50 22.87 14.50 23.23 855.5
IAC-11 42.50 68.50 115.00 32.50 14.50 4.00 13.00 13.00 6.50 4.00 3.50 26.78 11.50 28.32 533
PR-143 (14) 64.50 71.00 116.00 33.00 20.50 5.00 13.00 25.50 15.00 4.50 3.00 26.78 13.00 45.61 755
H18 46.50 75.50 116.00 41.50 10.00 4.00 15.50 16.50 9.50 6.50 4.50 32.29 12.00 16.19 867.5
PR-160-6 72.50 73.00 115.50 34.00 10.00 5.00 12.00 30.50 17.00 5.50 3.00 25.67 15.50 29.76 684.5
JSL 1 39.50 67.50 98.50 24.00 18.50 4.75 13.00 11.50 4.50 4.50 3.50 27.28 12.00 33.21 871
G9945 42.50 67.50 116.50 32.00 19.00 5.00 14.00 13.50 6.00 4.00 3.50 28.57 15.00 44.85 909.5
Hardee-1 47.50 68.50 95.00 37.00 18.00 4.25 13.00 11.50 4.00 5.50 3.50 26.78 14.50 41.25 1076
88
G01892 43.00 72.50 117.00 26.00 14.00 4.00 12.00 12.50 5.00 4.00 3.00 26.17 9.50 25 601
H3 44.00 67.00 99.50 23.50 19.50 5.00 13.00 10.50 4.50 4.50 3.50 26.97 11.50 33.14 1284.5
IAC-73-5115 66.00 76.00 116.50 52.00 13.50 4.00 25.50 46.50 24.00 8.00 3.00 11.80 17.00 18.05 938.5
PR-149-81-EP 42.00 69.50 115.50 25.00 12.00 4.00 13.50 17.00 8.50 4.50 3.00 25.83 12.00 22.18 871
AGS 214 46.00 75.00 98.00 26.50 25.50 4.00 14.00 13.50 6.00 5.00 3.50 24.87 11.50 38.17 1155
AGS-3-1 42.50 73.00 98.50 22.00 17.50 4.00 15.50 16.50 8.50 4.00 5.00 32.27 10.50 28.80 718.5
F81-7636-4 37.00 68.00 99.50 29.00 12.50 5.00 14.00 20.50 14.00 4.50 3.50 24.87 16.00 29.15 867
G00391 42.00 69.50 115.00 40.00 12.50 5.00 14.50 10.50 5.50 3.50 3.00 20.71 16.00 28.72 1126.5
G03705 48.50 75.00 116.00 52.00 19.50 5.00 13.00 25.50 12.50 4.00 3.00 22.71 9.50 30.00 929.5
G01853 47.50 72.00 115.50 32.50 12.50 5.00 14.00 21.50 17.50 4.50 3.00 21.54 15.50 30.38 664.5
H10 40.00 68.00 116.00 47.00 11.50 5.00 14.50 7.50 3.50 4.00 3.50 27.62 10.50 17.36 446
V1-1 42.00 67.50 116.00 25.00 15.50 5.00 14.00 11.50 5.00 4.50 3.00 21.55 11.50 27.54 819
AGS-299-2 47.50 69.00 116.00 34.00 22.50 5.00 14.50 12.50 6.00 4.50 3.50 27.62 12.00 37.26 955.5
PR-145-2 46.50 76.50 116.00 31.50 12.50 3.75 12.50 16.50 8.50 5.00 3.00 24.04 14.50 29.00 1026.5
FB1-7636 41.50 65.50 115.50 21.00 16.50 4.00 11.50 6.50 3.50 4.50 3.00 26.13 11.50 37.42 945.5
F82-7629-2 48.50 72.50 115.50 46.00 13.50 4.00 13.00 26.50 13.00 4.00 3.00 23.71 11.00 22.83 740.5
G00386 40.00 68.50 117.00 36.00 12.50 5.00 14.00 11.50 4.50 4.50 3.50 24.87 20.00 37.90 514.5
AGS-3 41.50 69.50 116.50 38.00 15.50 4.00 12.00 6.50 3.50 4.50 3.00 26.17 11.00 28.40 1209
G00141 41.50 72.50 116.50 46.50 14.50 4.50 13.50 16.50 9.50 4.00 3.00 22.25 13.00 28.78 517.5
H5 42.00 66.50 99.00 25.00 15.50 5.00 14.50 14.50 9.00 4.50 3.00 20.71 19.00 49.75 1177
Assosa local 63.50 102.00 117.00 40.50 16.50 4.50 13.50 15.50 8.00 4.00 3.50 22.25 16.50 39.61 1025
check-1
Mean 50.06 72.60 111.10 35.27 15.98 4.49 14.02 15.37 7.79 4.96 3.45 25.50 13.68 32.79 888.84
CV 1.90 1.11 0.79 3.36 5.73 5.80 9.48 6.50 13.76 11.86 13.50 7.59 7.69 4.17 3.43
LSD (5%) 1.96 1.62 1.75 2.45 1.84 0.53 2.71 2.05 2.25 1.21 0.91 3.85 2.17 2.90 62.91
89
Appendix figureIDendrogram of 49 soybean genotypes based on evaluation for 15 characters
grown at Jimma (2010/11).
90
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