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Seed Yield Stability and Genotype x Environment Interaction of Common Bean (Phaseolus vulgaris L.) Lines in Ethiopia
IJPBCS
Seed Yield Stability and Genotype x Environment
Interaction of Common Bean (Phaseolus vulgaris L.)
Lines in Ethiopia
Zeleke Ashango1
, Berhanu Amsalu2
, Kidane Tumisa3
, Kassaye Negash4
, Asnake Fikre5
1,2,3,4
Ethiopian Institute of Agricultural Research (EIAR); Melkassa Agricultural Research Center, P.O.Box 436, Adama,
Ethiopia.
5
Ethiopian Institute of Agricultural Research (EIAR); Addis Ababa, Ethiopia.
1*
Corresponding author: Zeleke Ashango*(zelekemarc2014@gmail.com)
When genotypes are introduced into a new and diverse production environments, occurrence of
significant genotype by environment interaction (GEI) complicates selection of stable
genotypes. Therefore, fifteen introduced and one check small red common bean lines were
evaluated at five representative dry bean growing locations of Ethiopia for seed yield
performance using a 4x4 triple lattice design in the 2013 and 2014 main cropping seasons to
estimate the magnitude of GEI effects and to identify broadly or specifically adapted lines.
Combined analysis of variance, Additive Main effects and Multiplicative Interaction (AMMI) and
Genotype plus Genotype x Environment interaction (GGE) biplot models were used to interpret
the data. Both the main and interaction effects were highly significant (p< 0.01) and
environment, line, and GEI explained 81.06%, 3.21% and 15.73% of variations, respectively,
indicating greater influence of environments and importance of simultaneous consideration of
mean performance and stability. PC1 and PC2 were highly significant (p < 0.01) and together
contributed nearly 60% variation in the GEI sum of squares. AMMI 1, GGE ranking, and GGE
comparison biplots enabled identification of both high seed yielding and broadly adapted lines,
KG-71-1, KG-71-23, and KG-71-44. Polygonal GGE biplot analysis enabled identification of four
mega-environments and specifically adapted lines. However, the specific adaptability of lines
was not repeated over years and thus, GEI couldn't be exploited and therefore, broadly adapted
lines were recommended for verification and release.
Key Words: AMMI, GGE, broad adaptation, line, environment, seed yield
INTRODUCTION
Common bean (Phaseolus vulgaris L.) is the most
important grain legume in nearly all lowland and mid
altitude areas of Ethiopia. It is produced primarily by
smallholder farmers both for cash and consumption. In
2014, it was cultivated by 3.34 million smallholders on
340 thousand hectare of land which is about 20% of total
farm land allocated for pulses (CSA, 2014). Its fastest
ripening at the critical food deficit period earlier than other
crops made it an ideal food deficit filler crop. Its suitability
for double or triple production per year enabled its
production on offseason free lands and relatively cheaper
labor force. Its reasonable protein content (22%) made it
the poor man's meat securing more than 16.7 million rural
people against hidden hunger. Despite its multifaceted
importance, most small red common bean cultivars at
production are more than 15 years old and are low seed
yielding. Therefore, advanced small red common bean
lines reported for their higher seed
International Journal of Plant Breeding and Crop Science
Vol. 3(2), pp. 135-144, October, 2016. © www.premierpublishers.org. ISSN: 2167-0449
Research Article
Seed Yield Stability and Genotype x Environment Interaction of Common Bean (Phaseolus vulgaris L.) Lines in Ethiopia
Ashango et al. 135
yield potential have been part of the research system
engagement.
When genotypes are performance tested at several
environments, the rankings usually differ as specified
difference in environment may produce different effect on
specific genotypes. Such inconsistent performance of
genotypes across environments is called genotype x
environment interaction (Asfaw et al., 2009). In highly
diverse environments there would likely be genotype x
environment interaction (GEI) is expected. Consequently,
it is not only average performance that is important in
selection of superior genotypes, but also the magnitude
of the interaction (Ebdon and Gauch, 2002b, Gauch and
Zobel, 1997) matters. GEI reflects differences in
adaptation and can be exploited by selecting for specific
adaptation or minimized by selecting for broad adaptation
(Adjei et al., 2010). These objectives can be achieved by
stratifying environments and by selecting adaptable
genotypes for each mega-environment or for broader
region. (Annicchairico, 2002). Multi-location evaluation of
genotypes provides useful information for this broader or
specific recommendation (Crossa, 1990). Mekbib (2003),
Asfaw et al. (2008), and Tsegaye et al. (2012) studied
seed yield performance of common bean cultivars
varying in growth habit and seed size at different parts of
Ethiopia and reported occurrence of significant GEI and
diversity of environments and cultivars.
Several biometrical methods had been developed and
used to analyze GEI, stability, and adaptability. In this
manuscript, AMMI and GGE models are considered for
multi-environment trials data analysis because they are
very useful tools to understand complex GEI, which
genotype outsmart where pattern discovery, and gaining
accuracy of yield estimates (Samonte et al., 2005;
Gauch, 2006; Yan et al., 2007; Gauch et al., 2008; Asfaw
et al., 2009; Namaratu et al., 2009). These approaches
integrate statistical and graphical analysis tools and help
the agronomists and plant breeders effectively perform
stability analysis (Fernandez, 2001). They are also free of
stringent statistical assumptions such as linearity of the
response of genotypes to a change in environments and
homogeneity of error variances among environments
(Anncchiarico, 1997).
The AMMI model is a hybrid analysis that incorporates
both the additive and multiplicative components of the
two-way data structure (Shafii et al. 1992; Shafii and
Price 1998). The AMMI biplot analysis is considered to be
an effective tool to diagnose the GEI patterns graphically.
In AMMI, the additive portion is separated from
interaction by analysis of variance (ANOVA). Then the
principal components analysis (PCA), which provides a
multiplicative model (Gabriel 1971; Zobel et al., 1988), is
applied to analyze the interaction effect from the additive
ANOVA model. The biplot display of PCA scores plotted
against each other provides visual inspection and
interpretation of the GEI components. Integrating biplot
display and genotypic stability statistics enables
genotypes to be grouped based on the similarity.
However, GGE biplot is superior to the AMMI graph in
mega-environment analysis (Yan et al., 2007).
As dry bean production environment of Ethiopia are
variable and test lines were new introductions from
International Center for Tropical Agriculture (CIAT), it is
necessary to evaluate lines before recommendation to
production agro-ecologies. Therefore, this research was
conducted to estimate the magnitude of line by
environment interaction effects and to analyze the
adaptability and stability of small red common bean lines
for seed yield performance in Ethiopia.
MATERIALS AND METHODS
Field experiment was conducted during the 2013 and
2014 main cropping seasons from July to October at five
representative locations of the dry bean growing agro-
ecologies of Ethiopia. The locations were namely
Melkassa, Alem Tena, Arsinegelle, Haramaya, and
Miesso. They were Abbreviated as MLK14= Melkassa
2014, MLK13 = Melkassa 2013, ALT14 = Alem Tena
2014, ALT13 = Alem Tena 2013, MIS14 = Miesso 2014,
MIS13 = Miesso 2013, ARN14 = Arsinegelle 2014,
ARN13 = Arsinegelle 2013, HRM14 = Haramaya 2014,
and HRM13 = Haramaya 2013. They were agricultural
research centers and sub centers. Experimental
materials were 15 small red common bean lines
introduced from CIAT through Pan African Bean
Research Alliance (PABRA) and one check. They were
coded as G1=KG-71-1, G2=KG-71-23, G3=KG-71-13,
G4=KG-71-20, G5=KG-71-44, G6=KG-71-26,
G7=SARBYT-2, G8=KG-67-11, G9=KG-103-11,
G10=F10 Black Sel New Bilfa-45, G11= F10 Black Sel
New Bilfa-46, G12=KG-71-21, G13=KG-71-46, G14=DAB
11, G15=SRE 194, G16= Dicta 105(check).
The experimental design used was a 4x4 triple lattice.
The plot size was 2.4m x 4m (9.6m
2
) with six rows of
spacing 40cm between rows and 10cm between plants.
The net harvested area was 6.4m2
, the central four rows.
Two seeds per hill were sown on rows with manual
drilling to ensure germination and good stands of the
common bean lines and then were thinned to one plant
per hill 12 days after emergence to achieve 480 plants
per plot. No fertilizer was applied as this is common
practice of most smallholder farmers for common bean
cultivars and other cultural practices like seed bed
preparation and weeding were followed as per
recommendations for released small red common bean
cultivars.
Seed yield data were collected from the central four rows
of the plot and adjusted to 14% seed moisture
Seed Yield Stability and Genotype x Environment Interaction of Common Bean (Phaseolus vulgaris L.) Lines in Ethiopia
Int. J. Plant Breeding Crop Sci. 136
using the equation (Hong and Ellis, 1996) Yadj =
100−MC
100−14
∗ Y , where: 𝑌𝑎𝑑𝑗 was moisture adjusted yield, Y
was unadjusted yield, and MC was measured moisture
content (%).
Combined ANOVA over locations and years and AMMI
model analysis of variance were done using Genstat
version 17 statistical software package. Data were
combined after Bartlett's test for homogeneity of error
variance. The combined ANOVA model of fixed effects
lines and random effects locations, years, and
interactions was:
𝑦𝑖𝑗𝑘 = 𝜇 + 𝐺𝑖 + 𝐿𝑗 + 𝑌𝑘 + (𝐺𝐿)𝑖𝑗 +
(𝐺𝑌)𝑖𝑘 + (𝐿𝑌)𝑗𝑘 + (𝐺𝐿𝑌)𝑖𝑗𝑘 +∈𝑖𝑗𝑘 Where: yijk is the
mean yield across replicates of the ith
line in the
jth
location and kth
year, µ is the grand mean, Gi is the
additive effect of ith
line, Lj is the additive effect of jth
location, Yk is the additive effect of kth
year, (𝐺𝐿)𝑖𝑗 ,
(𝐺𝑌)𝑖𝑘, (𝐿𝑌)𝑗𝑘 , and (𝐺𝐿𝑌)𝑖𝑗𝑘 are the interaction effects of
lines, locations and years. ∈𝑖𝑗𝑘 is the error assumed to be
normally and independently distributed as (0 , σ2
r)
where σ2
is the pooled error variance and r is the
number of replicates.
Through AMMI model, GEI was further partitioned into
IPCA components and the AMMI model (Zobel et al.,
1988) used was:
Where: Yij is the mean yield across replicates of the ith
cultivar in the jth
environment, µ is the grand mean, Gi is
the additive effect of ith
line, Ej is the additive effect of jth
environment, Kn is the singular value of the IPCA axis n,
Uin and Sjn are scores of line i and environment j for the
IPC axis n, respectively, Qij is residual for the first n
multiplicative components, and ɛij. is the residual error
assumed to be normally and independently distributed as
(0 , σ2
r) (where σ2
is the pooled error variance and r
is the number of replicates).
To show a clear insight into specific lines by environment
interaction (GEI) combinations and the general pattern of
adaptation, biplots of lines and environments (AMMI1,
AMMI2, and different GGE) were developed using
Genstat version17 statistical software package. The
predicted absolute IPCA 1 score values for ranking of
lines based on adaptability in Figure 1 were taken from
AMMI model analysis of variance to aid visual inspection
of the position of each line on the graph. The different
GGE biplots from Figure 3 - 8 were generated using
similar data, but varying scaling for lines and
environments.
RESULTS AND DISCUSSION
Analysis of GEI
The mean squares from the combined analysis of
variance over locations and years from both combined
ANOVA and AMMI models are presented in Table 2. The
analysis showed that lines (G), locations (L), years (Y),
lines x location (GL), lines x year (LY), location x year
(LY), and lines x location x year (GLY) effects were highly
significant (p < 0.01). This indicated the diversity of
locations and years and presence of substantial genetic
differences among the lines for seed yield performance.
Similar findings were reported by Mekbib (2003), Asfaw
et al. (2008) and Tamene and Tadese (2014) for common
bean cultivars performance and their growing
environments in Ethiopia. The significant GL, GY, LY,
and GLY were also indicated that the relative
performance of lines at different locations and years was
not similar.
Combined analysis of variance partitions the variation in
a two way factorial multi-environment trial data into
genotype main effects, environment main effects, and
genotype by environment interaction effects with the most
common outcome of largest environment main effects
followed by the interaction effects and then the variety
main effects (Gauch, 2006; Yan and Kang, 2003). In the
present study, the largest effects of environment
(81.06%) followed by GEI effects (15.73%) and then by
lines main effects (3.21%) were observed (Table 2). From
the portion of variation explained by environment,
location alone contributed 69.84% and years explained
0.53%. This indicated greater influence of spatial
variation than temporal variation on lines seed yield
performance. The GL, GY, LY, and GLY effects were
10.06%, 3.77%, 6.35%, and 6.40%, respectively. Hence,
GEI exerted five times larger effect than lines main effect
to the observed phenotype and highly significantly
complicated selection of superior and adaptable lines.
Therefore, simultaneous consideration of both high mean
seed yield performance (main effects) and GEI (stability)
is very important in selecting among the small red
common bean lines evaluated. This result is in
agreement with the reports of Mekbib (2003), Asfaw et
al. (2008), and Tsegaye et al. (2012).
Combined ANOVA determines if GEI is a significant
source of variation or not and estimates it, but does not
provide insight into the patterns of genotypes or
environments that give rise to the interaction (Samonte et
al., 2005). Therefore, the combined data was also
analyzed using AMMI model that further partitions GEI
into interaction principal component axis (IPCA)
components. Hence, the AMMI model analysis had
partitioned the GEI into the first two significant IPCAs
with contributions of IPCA1 (39.45%) and IPCA2
(17.38%). The remaining residuals were not significant
(Table 2). Therefore, IPCA1 and IPCA2 alone were
adequately predicted the variation in this data structure
and thus, the overall pattern of lines interaction with
environments was interpreted using AMMI1, AMMI2, and
GGE biplot models.
  ijij
n
nininjiij QSUkEGY   1
Seed Yield Stability and Genotype x Environment Interaction of Common Bean (Phaseolus vulgaris L.) Lines in Ethiopia
Ashango et al. 137
Table 1. Climatic characteristics of the test locations.
No Locations Abbreviation
Altitude
(m .a .s. l.)
Annual
rainfall (mm) Temperature (℃ )
Min. Max.
1 Melkassa MLK 1550 768 13.80 28.60
2 Alem Tena ALT 1660 832 13.50 27.50
3 Miesso MIS 1394 727 15.00 30.60
5 Arsinegelle ARN 1951 915 12.00 25.50
6 Haramaya HRM 1950 790 11.00 26.00
Table 2. Mean squares of combined ANOVA and AMMI models analysis of variance of small red common bean lines
evaluated at five locations of Ethiopia during the main cropping seasons in the 2013 and 2014.
Mean squares
Sources of variation Degree of freedom
Combined
ANOVA AMMI
Explained %
of treatments
SS
Blocks 2 99628
ns
Blocks (E) 20 787107
**
Treatments 159 2667359
**
Lines (G) 15 907918
*
907918
*
3.21
Location (L) 4 78462850
**
69.84
Year (Y) 1 2393364
**
0.53
GL 60 753755
*
10.06
GY 15 1130280
**
3.77
LY 4 7138233
**
6.35
GLY 60 479421
**
6.40
Environment (E) 9 38199744
**
81.06
GEI 135 494070
**
15.73
IPCA1 23 1144351
**
39.46
IPCA2 21 552136
**
17.38
Residual 91 316313
ns
Error 318(300) 281383 246457
* &**= significant at 0.05 and 0.01 probability levels, ns=Non significant, GEI=Genotype by environment interaction, GL=Genotype
by location interaction, GY=Genotype by year interaction, LY=Location by year interaction, GLY=Genotype by location by year
interaction, (300)=Degree of freedom for error term of AMMI model, SS=Sum of squares.
Stability Analysis
AMMI Biplots analysis
AMMI biplots are recently preferred biplots to visualize
adaptability and stability of genotypes over test
environments (Gauch and Zobel, 1988; Gauch and
Zobel, 1996; Gauch, 2006; Gauch et al., 2008). This is
because they are effective tools to diagnose the GEI
patterns graphically. In AMMI1 biplot, the genotypes
with IPCA1 scores close to zero express general
adaptation and the larger scores depict more specific
adaptation in combination with environments of the same
sign IPCA1 scores (Ebdon and Gauch, 2002a).
Furthermore, the relative magnitude and direction of
genotypes along the abscissa and ordinate axis in
biplot is also important to understand the response
pattern of genotypes across environments and to
Seed Yield Stability and Genotype x Environment Interaction of Common Bean (Phaseolus vulgaris L.) Lines in Ethiopia
Int. J. Plant Breeding Crop Sci. 138
Figure 1. AMMI 1 biplot showing mean performance and adaptability of small red common bean lines
over environments
differentiate high yielding and adaptable genotypes
(Samonte et al., 2005). Accordingly, in Figure 1, G2, G1,
and G5 placed relatively close to zero IPCA1 score line
and performed above the overall mean were generally
adapted to all small red common bean lines growing
environments of Ethiopia. The high yielding line (G8) had
similar sign IPCA1 scores with IPCA1 score sign of
HRM13 and ALT13 showed positive interaction with
these environments. Thus, it was specifically adapted to
these environments and similar agro-ecologies. Similarly,
the high yielding lines (G15 and G9) with similar sign of
IPCA1 score to MLK13, MLK14, and HRM14 showed
positive interaction with these environments and
specifically adapted to these and similar small red
common bean cultivars growing environments of
Ethiopia.
Considering environments, MLK14, MLK13 and HRM14
exhibited high seed yield performance (Fig. 1). Thus, they
are better environments for commercial production of
common bean lines found specifically or widely adapted
to them. MIS13, and MIS14 were low seed yield potential
environments. As it is located furthest away from zero
line of IPCA1 score, MLK14 showed greatest interaction
with lines. This could be due to the less fertile soils of
Miesso and better moisture of Melkassa and Haramaya
(Table 1).
In overall, the lines adaptability/stability ranking for seed
yield performance based on lower absolute IPCA1 score
value which measures the distance of each line from zero
value line, i.e., absolute adaptability line, was G13 (0.33)
> G4(1.04) > G5 (2.37) > G2 (3.01) > G3(4.27) > G11
(4.67) > G1(6.16) > G14 (7.46) > G12 (9.17) > G10 (9.44)
> G8 (11.28) > G9(11.56 ) > G7(136.91) >G15 (17.86) >
G16 (39.29) (Fig. 1). This indicated that G13 was the
most stable small red common bean line , but below
average seed yielding and adapted to low seed yielding
location Miesso and similar agro-ecologies while G16
was the most unstable, low seed yielding, and un-
adapted to Ethiopa compared to the lines in the test set.
The AMMI1 biplot, Fig. 1, had visualized not only the
lines performance in relation to adaptability and mean
seed yield performance, but also revealed presence of
two mega-environments. Five low and medium seed yield
potential environments (MIS13, MIS14, ARN13, HRM13,
and ALT13) with similar IPCA1 scores had formed one
mega-environment whereas another five medium and
high seed yield potential environments (ALT14, ARN14,
MLK14, HRM14, and MLK13) had formed another mega-
environment. However, its mega-environment
classification is more general and didn’t show detailed
specific adaptation of lines (Yan et al., 2007). Therefore,
more specific adaptability of lines was explored using
AMMI 2 biplot (Fig. 2).
In AMMI 2 biplot, Figure 2, the small red common bean
lines (G16, G5, G13, G15, and G7) placed furthest away
from the biplot origin expressed a highly interactive
behavior (positively or negatively) whereas G3 and G4
placed relatively close to the biplot origin expressed less
interaction and more adapted to all environments. This is
because, in AMMI 2 biplot, the distances from the biplot
origin are indicative of the amount of interaction
exhibited by genotypes over environments or
environments over genotypes and genotypes located
near the biplot origin are less responsive than the vertex
genotypes indicating general adaptability to all
environments (Voltas et al.,2002). Environments, MLK 14
and MIS 14, with longer vectors were very interactive and
discriminated the differences among genotypes more
than other environments with shorter vectors, which were
less interactive and provided little information about the
Seed Yield Stability and Genotype x Environment Interaction of Common Bean (Phaseolus vulgaris L.) Lines in Ethiopia
Ashango et al. 139
Figure 2. AMMI 2 biplot showing general and specific adaptability of small red common bean lines
over environments
Figure 3. Genotype focusing scaled GGE scatter biplot showing mean performance and stability of
small red common bean lines.
differences among the lines' seed yield performances as
reported by (Yan, 2002).
Similarly, in Figure 2, the angles between genotype,
environment, or between genotype and environment
vectors determine the nature of GEI. The interaction is
positive for acute angles, zero for right angles, and
negative for obtuse angles (Kandus et al., 2010).
Accordingly, G8, G11, and G5 which made acute angles
with ARN13 and ALT13 vector showed positive
interaction and specifically adapted to these
environments. The lines, G15, G1, G2, G5, made acute
angles with ALT14 and MLK14 vectors and interacted
highly positively with them , therefore, were specifically
adapted to them. G9, G7, and G10 were specifically
adapted to MLK13. Again, G7, G9 and G6 interacted
highly positively with ARN14 and HRM14 and specifically
adapted to them. The low seed yielding line, G13, had
made acute angle with low seed yield potential
environments, MIS13 and MIS14, vectors was specifically
best line for them. MIS13 being placed close to the biplot
origin showed less interaction with lines than other
environments and provided little information about lines'
seed yield performance. The low seed yielding line, G16,
being placed furthest away from all environments was un-
adapted to all environments compared to the lines in the
test set.
GGE Biplots Analysis
Mean Seed Yield Performance and Stability of Lines
In GGE scatter biplot scaled focusing genotype,
PC1score estimates mean yield with its zero line
indicating average performance while the least absolute
PC2 score shows top stability (Yan and Kang, 2002).
Thus, in Figure 3, lines which had PC1 scores greater
Seed Yield Stability and Genotype x Environment Interaction of Common Bean (Phaseolus vulgaris L.) Lines in Ethiopia
Int. J. Plant Breeding Crop Sci. 140
Figure 4. Average environment coordination (AEC) view of the GGE comparison biplot showing
performance of lines compared with best line performance.
than zero viz, G15, G1, G9, G6, G10, G7, G2, and G5
were higher seed yielding lines while lines with PC1
scores less than zero, G8, G11, G3, G14, G13, G4, G12,
and G14 were lower seed yielding lines. G14 placed
closest to zero line of PC2 score was the most stable, but
low seed yielding. Lines placed furthest away from zero
PC2 line were unstable.
In genotype focusing scaled GGE ranking biplot, Figure
5, lines, G16, G14, G4, G13, G7, G10, and G12, which
had fallen below the AEC ordinate showed below
average seed yield performance whereas lines, G15,
G1, G6, G9, G2, G5, G11, G8, and G3, which had fallen
above the average environment coordination (AEC)
ordinate performed above average. G1, G2, G3, G11,
and G5 which had performed above average and had
relatively shortest projection vectors from AEC line were
both high seed yielding and widely adapted. On the other
hand G4, G13, and G14 which had fallen below AEC
ordinate axis and with shortest projection vectors from
AEC line were widely adapted, but low seed yielding.
This is because, in genotype focusing scaled ranking
GGE biplot, AEC approximates the genotypes' main
effect, that is, mean performance with the arrow pointing
to greater genotype main effect while its ordinate
approximates GEI effects, that is, stability with increasing
GEI effects and instability away from the origin at both
directions (Yan et al., 2001; Yan and Hunt, 2002).
In genotype focusing scaled GGE comparison biplot,
Figure 4, line, G1, is ideal line and G5, G2, G15, G11,
and G8 placed relatively closest to G1 were both higher
seed yielding and widely adapted lines compared to other
lines. In genotype focusing scaled comparison GGE
biplot, a genotype located nearest to the central
concentric circle is both high seed yielding and most
stable. It is considered as an ideal genotype and
genotypes fall closest to it are also considered as
desirable (Yan, 2002).
In symmetrically scaled polygonal GGE biplot, Figure 6,
seven sectors of which four with environments were
observed. MLK14, ALT14, HRM14, ARN14, and MIS13
were clustered in one sector and could be considered as
one mega-environment for small red common bean lines
evaluation and recommendation. Their higher seed
yielding lines were G15, G1, G5, and G2. ALT13 and
ARN13 had entered into one sectors had also formed
another mega-environment with their higher seed yielding
lines G8, and G11. MLK13 alone made one mega-
environment and its higher seed yielding lines were G7
and G10. MIS13 and HRN13 had made fourth mega-
environment and their winning lines were G12 and G3.
These specific adaptability and mega-environments
interpretations are because, in symmetrically scaled
polygon view of GGE biplot, connecting the extreme
genotypes forms a polygon and the perpendicular lines to
the sides of the polygon form sectors of genotypes and
environments, which are considered as separate mega-
environments (Gauch and Zobel, 1997; Yan et al., 2000).
GEI reflects differences in adaptation and can be
exploited by selecting for specific adaptation if the trend
in specific adaptability of genotypes is repeatable over
years (Annicchiarico, 2002; Yan et al., 2007). However, in
this study, the specific adaptability trend was not
repeated over years as different environments were
grouped differently in two years (Figures 7). Therefore,
GEI couldn't be exploited and should be minimized by
selecting for broad adaptation. Thus, broadly adapted
lines, G1, G2, and G5, were recommended for
verification and release.
Seed Yield Stability and Genotype x Environment Interaction of Common Bean (Phaseolus vulgaris L.) Lines in Ethiopia
Ashango et al. 141
Figure 5. Average environment coordination (AEC) view of the GGE ranking biplot showing mean
performance and stability of lines.
Discriminating ability, Representativeness, and
Relationships of test Environments
In environment focusing scaled vector view of GGE
biplot, the cosine of the angles between environment
vectors show relationships between test environments
with acute angles indicating strong correlation, obtuse
angles strong negative correlation or cross over GEI of
genotypes, and right angle showing no correlation (Yan
and Tinker, 2006). Hence, in Figure 8 right, (ALT13 and
ARN13) and (MLK14 and ALT14) with acute angles
between them were strongly correlated and indicated
significant influence of years on genotypes' seed yield
performance. Thus, similar information could be obtained
by dropping either of one environment for small red
common bean lines evaluation by reducing cost of multi-
location replicating trials. Therefore, it is better to drop
Alem Tena from trial sites as similar information can be
obtained from Melkassa and Arsinegelle. MLK14 with the
longest vector length was the most discriminating and more
Figure 6. Symmetrically scaled polygonal GGE biplot showing specific adaptability of lines.
Seed Yield Stability and Genotype x Environment Interaction of Common Bean (Phaseolus vulgaris L.) Lines in Ethiopia
Int. J. Plant Breeding Crop Sci. 142
Figure 7. Symmetrically scaled polygonal GGE biplots showing specific adaptability of lines during 2013 (right) and 2014
(left).
Figures 8. Environment focusing scaled vector view of GGE biplot (right) and average environment coordination (AEC)
view of GGE biplot (left) showing relationships among test environments.
informative environment while MIS13 with the shortest
vector was the least discriminating and less informative
environment. The lines interaction with environments was
crossover in MLK14 and MIS14 as indicated by obtuse
angle between their vectors (Fig 8, right).
Similarly, in environment focusing scaled comparison
GGE biplot (Fig. 8 left), a test environment with smallest
angle between the AEC abscissa is the most
representative (Yan and Tinker, 2006). Hence, MLK14
laid on the AEC abscissa line was the most
Seed Yield Stability and Genotype x Environment Interaction of Common Bean (Phaseolus vulgaris L.) Lines in Ethiopia
Ashango et al. 143
representative of all environments followed by ALT14.
Therefore, MLK14 was both most representative and
most discriminating. This is because Melkassa is nearly
optimum environment in terms of all edaphic, climatic,
and biotic seed yield limiting factors, but Miesso is
characterized by less fertile soils, low moisture
availability, and higher temperature. Alem Tena,
Arsinegelle, and Haramaya are average on all seed
yield limiting factors and are medium seed yielding
environments.
CONCLUSION
GEI is differential phenotypic performance of genetically
uniform genotypes across test environments. It occurs
because different genotypes have varying genetic
potentials to adjust themselves to variable environments.
Small red common bean lines evaluated by this study
had highly significant genetic differences for seed yield
performance across environments. Spatial variation in
environments was more profound than temporal
variations in exerting effects on lines' seed yield
performance. AMMI 1, GGE ranking, and GGE
comparison biplots enabled identification of broadly
adapted and high seed yielding lines better than AMMI 2
and polygonal GGE biplots. Small red common bean
lines, KG-71-1, KG-71-23, and KG-71-44 were both high
seed yielding and broadly adapted to dry bean growing
environments of Ethiopia. AMMI 2 and polygonal GGE
biplots enabled selection of specifically adapted lines, but
the specific adaptability show of polygonal GGE biplot is
clearer than AMMI 1 and AMMI 2 biplots. Environment
focusing scaled GGE biplots also enabled identification of
ideal (Melkassa), least informative (Miesso), and
redundant (Alem Tena) environments for small red
common bean lines evaluation. However, the specific
adaptability of lines was not consistent over years.
Hence, GEI couldn't be exploited and broadly adapted
lines, KG-71-1, KG-71-23, and KG-71-44, were
recommended for verification and release.
ACKNOWLEDGEMENT
The authors thank MARC and CIAT-PABRA for financial
support and experimental materials provision. Our
thanks are also extended to those all who participated
during execution of the experiment.
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Citation: Ashango Z, Amsalu B, Fikre A, Tumisa K,
Negash K (2016). Seed Yield Stability and Genotype x
Environment Interaction of Common Bean (Phaseolus
vulgaris L.) Lines in Ethiopia. International Journal of
Plant Breeding and Crop Science, 3(2): 135-144.
Copyright: © 2016 Ashango et al. This is an open-
access article distributed under the terms of the Creative
Commons Attribution License, which permits unrestricted
use, distribution, and reproduction in any medium,
provided the original author and source are cited.

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Seed Yield Stability and Genotype x Environment Interaction of Common Bean (Phaseolus vulgaris L.) Lines in Ethiopia

  • 1. Seed Yield Stability and Genotype x Environment Interaction of Common Bean (Phaseolus vulgaris L.) Lines in Ethiopia IJPBCS Seed Yield Stability and Genotype x Environment Interaction of Common Bean (Phaseolus vulgaris L.) Lines in Ethiopia Zeleke Ashango1 , Berhanu Amsalu2 , Kidane Tumisa3 , Kassaye Negash4 , Asnake Fikre5 1,2,3,4 Ethiopian Institute of Agricultural Research (EIAR); Melkassa Agricultural Research Center, P.O.Box 436, Adama, Ethiopia. 5 Ethiopian Institute of Agricultural Research (EIAR); Addis Ababa, Ethiopia. 1* Corresponding author: Zeleke Ashango*(zelekemarc2014@gmail.com) When genotypes are introduced into a new and diverse production environments, occurrence of significant genotype by environment interaction (GEI) complicates selection of stable genotypes. Therefore, fifteen introduced and one check small red common bean lines were evaluated at five representative dry bean growing locations of Ethiopia for seed yield performance using a 4x4 triple lattice design in the 2013 and 2014 main cropping seasons to estimate the magnitude of GEI effects and to identify broadly or specifically adapted lines. Combined analysis of variance, Additive Main effects and Multiplicative Interaction (AMMI) and Genotype plus Genotype x Environment interaction (GGE) biplot models were used to interpret the data. Both the main and interaction effects were highly significant (p< 0.01) and environment, line, and GEI explained 81.06%, 3.21% and 15.73% of variations, respectively, indicating greater influence of environments and importance of simultaneous consideration of mean performance and stability. PC1 and PC2 were highly significant (p < 0.01) and together contributed nearly 60% variation in the GEI sum of squares. AMMI 1, GGE ranking, and GGE comparison biplots enabled identification of both high seed yielding and broadly adapted lines, KG-71-1, KG-71-23, and KG-71-44. Polygonal GGE biplot analysis enabled identification of four mega-environments and specifically adapted lines. However, the specific adaptability of lines was not repeated over years and thus, GEI couldn't be exploited and therefore, broadly adapted lines were recommended for verification and release. Key Words: AMMI, GGE, broad adaptation, line, environment, seed yield INTRODUCTION Common bean (Phaseolus vulgaris L.) is the most important grain legume in nearly all lowland and mid altitude areas of Ethiopia. It is produced primarily by smallholder farmers both for cash and consumption. In 2014, it was cultivated by 3.34 million smallholders on 340 thousand hectare of land which is about 20% of total farm land allocated for pulses (CSA, 2014). Its fastest ripening at the critical food deficit period earlier than other crops made it an ideal food deficit filler crop. Its suitability for double or triple production per year enabled its production on offseason free lands and relatively cheaper labor force. Its reasonable protein content (22%) made it the poor man's meat securing more than 16.7 million rural people against hidden hunger. Despite its multifaceted importance, most small red common bean cultivars at production are more than 15 years old and are low seed yielding. Therefore, advanced small red common bean lines reported for their higher seed International Journal of Plant Breeding and Crop Science Vol. 3(2), pp. 135-144, October, 2016. © www.premierpublishers.org. ISSN: 2167-0449 Research Article
  • 2. Seed Yield Stability and Genotype x Environment Interaction of Common Bean (Phaseolus vulgaris L.) Lines in Ethiopia Ashango et al. 135 yield potential have been part of the research system engagement. When genotypes are performance tested at several environments, the rankings usually differ as specified difference in environment may produce different effect on specific genotypes. Such inconsistent performance of genotypes across environments is called genotype x environment interaction (Asfaw et al., 2009). In highly diverse environments there would likely be genotype x environment interaction (GEI) is expected. Consequently, it is not only average performance that is important in selection of superior genotypes, but also the magnitude of the interaction (Ebdon and Gauch, 2002b, Gauch and Zobel, 1997) matters. GEI reflects differences in adaptation and can be exploited by selecting for specific adaptation or minimized by selecting for broad adaptation (Adjei et al., 2010). These objectives can be achieved by stratifying environments and by selecting adaptable genotypes for each mega-environment or for broader region. (Annicchairico, 2002). Multi-location evaluation of genotypes provides useful information for this broader or specific recommendation (Crossa, 1990). Mekbib (2003), Asfaw et al. (2008), and Tsegaye et al. (2012) studied seed yield performance of common bean cultivars varying in growth habit and seed size at different parts of Ethiopia and reported occurrence of significant GEI and diversity of environments and cultivars. Several biometrical methods had been developed and used to analyze GEI, stability, and adaptability. In this manuscript, AMMI and GGE models are considered for multi-environment trials data analysis because they are very useful tools to understand complex GEI, which genotype outsmart where pattern discovery, and gaining accuracy of yield estimates (Samonte et al., 2005; Gauch, 2006; Yan et al., 2007; Gauch et al., 2008; Asfaw et al., 2009; Namaratu et al., 2009). These approaches integrate statistical and graphical analysis tools and help the agronomists and plant breeders effectively perform stability analysis (Fernandez, 2001). They are also free of stringent statistical assumptions such as linearity of the response of genotypes to a change in environments and homogeneity of error variances among environments (Anncchiarico, 1997). The AMMI model is a hybrid analysis that incorporates both the additive and multiplicative components of the two-way data structure (Shafii et al. 1992; Shafii and Price 1998). The AMMI biplot analysis is considered to be an effective tool to diagnose the GEI patterns graphically. In AMMI, the additive portion is separated from interaction by analysis of variance (ANOVA). Then the principal components analysis (PCA), which provides a multiplicative model (Gabriel 1971; Zobel et al., 1988), is applied to analyze the interaction effect from the additive ANOVA model. The biplot display of PCA scores plotted against each other provides visual inspection and interpretation of the GEI components. Integrating biplot display and genotypic stability statistics enables genotypes to be grouped based on the similarity. However, GGE biplot is superior to the AMMI graph in mega-environment analysis (Yan et al., 2007). As dry bean production environment of Ethiopia are variable and test lines were new introductions from International Center for Tropical Agriculture (CIAT), it is necessary to evaluate lines before recommendation to production agro-ecologies. Therefore, this research was conducted to estimate the magnitude of line by environment interaction effects and to analyze the adaptability and stability of small red common bean lines for seed yield performance in Ethiopia. MATERIALS AND METHODS Field experiment was conducted during the 2013 and 2014 main cropping seasons from July to October at five representative locations of the dry bean growing agro- ecologies of Ethiopia. The locations were namely Melkassa, Alem Tena, Arsinegelle, Haramaya, and Miesso. They were Abbreviated as MLK14= Melkassa 2014, MLK13 = Melkassa 2013, ALT14 = Alem Tena 2014, ALT13 = Alem Tena 2013, MIS14 = Miesso 2014, MIS13 = Miesso 2013, ARN14 = Arsinegelle 2014, ARN13 = Arsinegelle 2013, HRM14 = Haramaya 2014, and HRM13 = Haramaya 2013. They were agricultural research centers and sub centers. Experimental materials were 15 small red common bean lines introduced from CIAT through Pan African Bean Research Alliance (PABRA) and one check. They were coded as G1=KG-71-1, G2=KG-71-23, G3=KG-71-13, G4=KG-71-20, G5=KG-71-44, G6=KG-71-26, G7=SARBYT-2, G8=KG-67-11, G9=KG-103-11, G10=F10 Black Sel New Bilfa-45, G11= F10 Black Sel New Bilfa-46, G12=KG-71-21, G13=KG-71-46, G14=DAB 11, G15=SRE 194, G16= Dicta 105(check). The experimental design used was a 4x4 triple lattice. The plot size was 2.4m x 4m (9.6m 2 ) with six rows of spacing 40cm between rows and 10cm between plants. The net harvested area was 6.4m2 , the central four rows. Two seeds per hill were sown on rows with manual drilling to ensure germination and good stands of the common bean lines and then were thinned to one plant per hill 12 days after emergence to achieve 480 plants per plot. No fertilizer was applied as this is common practice of most smallholder farmers for common bean cultivars and other cultural practices like seed bed preparation and weeding were followed as per recommendations for released small red common bean cultivars. Seed yield data were collected from the central four rows of the plot and adjusted to 14% seed moisture
  • 3. Seed Yield Stability and Genotype x Environment Interaction of Common Bean (Phaseolus vulgaris L.) Lines in Ethiopia Int. J. Plant Breeding Crop Sci. 136 using the equation (Hong and Ellis, 1996) Yadj = 100−MC 100−14 ∗ Y , where: 𝑌𝑎𝑑𝑗 was moisture adjusted yield, Y was unadjusted yield, and MC was measured moisture content (%). Combined ANOVA over locations and years and AMMI model analysis of variance were done using Genstat version 17 statistical software package. Data were combined after Bartlett's test for homogeneity of error variance. The combined ANOVA model of fixed effects lines and random effects locations, years, and interactions was: 𝑦𝑖𝑗𝑘 = 𝜇 + 𝐺𝑖 + 𝐿𝑗 + 𝑌𝑘 + (𝐺𝐿)𝑖𝑗 + (𝐺𝑌)𝑖𝑘 + (𝐿𝑌)𝑗𝑘 + (𝐺𝐿𝑌)𝑖𝑗𝑘 +∈𝑖𝑗𝑘 Where: yijk is the mean yield across replicates of the ith line in the jth location and kth year, µ is the grand mean, Gi is the additive effect of ith line, Lj is the additive effect of jth location, Yk is the additive effect of kth year, (𝐺𝐿)𝑖𝑗 , (𝐺𝑌)𝑖𝑘, (𝐿𝑌)𝑗𝑘 , and (𝐺𝐿𝑌)𝑖𝑗𝑘 are the interaction effects of lines, locations and years. ∈𝑖𝑗𝑘 is the error assumed to be normally and independently distributed as (0 , σ2 r) where σ2 is the pooled error variance and r is the number of replicates. Through AMMI model, GEI was further partitioned into IPCA components and the AMMI model (Zobel et al., 1988) used was: Where: Yij is the mean yield across replicates of the ith cultivar in the jth environment, µ is the grand mean, Gi is the additive effect of ith line, Ej is the additive effect of jth environment, Kn is the singular value of the IPCA axis n, Uin and Sjn are scores of line i and environment j for the IPC axis n, respectively, Qij is residual for the first n multiplicative components, and ɛij. is the residual error assumed to be normally and independently distributed as (0 , σ2 r) (where σ2 is the pooled error variance and r is the number of replicates). To show a clear insight into specific lines by environment interaction (GEI) combinations and the general pattern of adaptation, biplots of lines and environments (AMMI1, AMMI2, and different GGE) were developed using Genstat version17 statistical software package. The predicted absolute IPCA 1 score values for ranking of lines based on adaptability in Figure 1 were taken from AMMI model analysis of variance to aid visual inspection of the position of each line on the graph. The different GGE biplots from Figure 3 - 8 were generated using similar data, but varying scaling for lines and environments. RESULTS AND DISCUSSION Analysis of GEI The mean squares from the combined analysis of variance over locations and years from both combined ANOVA and AMMI models are presented in Table 2. The analysis showed that lines (G), locations (L), years (Y), lines x location (GL), lines x year (LY), location x year (LY), and lines x location x year (GLY) effects were highly significant (p < 0.01). This indicated the diversity of locations and years and presence of substantial genetic differences among the lines for seed yield performance. Similar findings were reported by Mekbib (2003), Asfaw et al. (2008) and Tamene and Tadese (2014) for common bean cultivars performance and their growing environments in Ethiopia. The significant GL, GY, LY, and GLY were also indicated that the relative performance of lines at different locations and years was not similar. Combined analysis of variance partitions the variation in a two way factorial multi-environment trial data into genotype main effects, environment main effects, and genotype by environment interaction effects with the most common outcome of largest environment main effects followed by the interaction effects and then the variety main effects (Gauch, 2006; Yan and Kang, 2003). In the present study, the largest effects of environment (81.06%) followed by GEI effects (15.73%) and then by lines main effects (3.21%) were observed (Table 2). From the portion of variation explained by environment, location alone contributed 69.84% and years explained 0.53%. This indicated greater influence of spatial variation than temporal variation on lines seed yield performance. The GL, GY, LY, and GLY effects were 10.06%, 3.77%, 6.35%, and 6.40%, respectively. Hence, GEI exerted five times larger effect than lines main effect to the observed phenotype and highly significantly complicated selection of superior and adaptable lines. Therefore, simultaneous consideration of both high mean seed yield performance (main effects) and GEI (stability) is very important in selecting among the small red common bean lines evaluated. This result is in agreement with the reports of Mekbib (2003), Asfaw et al. (2008), and Tsegaye et al. (2012). Combined ANOVA determines if GEI is a significant source of variation or not and estimates it, but does not provide insight into the patterns of genotypes or environments that give rise to the interaction (Samonte et al., 2005). Therefore, the combined data was also analyzed using AMMI model that further partitions GEI into interaction principal component axis (IPCA) components. Hence, the AMMI model analysis had partitioned the GEI into the first two significant IPCAs with contributions of IPCA1 (39.45%) and IPCA2 (17.38%). The remaining residuals were not significant (Table 2). Therefore, IPCA1 and IPCA2 alone were adequately predicted the variation in this data structure and thus, the overall pattern of lines interaction with environments was interpreted using AMMI1, AMMI2, and GGE biplot models.   ijij n nininjiij QSUkEGY   1
  • 4. Seed Yield Stability and Genotype x Environment Interaction of Common Bean (Phaseolus vulgaris L.) Lines in Ethiopia Ashango et al. 137 Table 1. Climatic characteristics of the test locations. No Locations Abbreviation Altitude (m .a .s. l.) Annual rainfall (mm) Temperature (℃ ) Min. Max. 1 Melkassa MLK 1550 768 13.80 28.60 2 Alem Tena ALT 1660 832 13.50 27.50 3 Miesso MIS 1394 727 15.00 30.60 5 Arsinegelle ARN 1951 915 12.00 25.50 6 Haramaya HRM 1950 790 11.00 26.00 Table 2. Mean squares of combined ANOVA and AMMI models analysis of variance of small red common bean lines evaluated at five locations of Ethiopia during the main cropping seasons in the 2013 and 2014. Mean squares Sources of variation Degree of freedom Combined ANOVA AMMI Explained % of treatments SS Blocks 2 99628 ns Blocks (E) 20 787107 ** Treatments 159 2667359 ** Lines (G) 15 907918 * 907918 * 3.21 Location (L) 4 78462850 ** 69.84 Year (Y) 1 2393364 ** 0.53 GL 60 753755 * 10.06 GY 15 1130280 ** 3.77 LY 4 7138233 ** 6.35 GLY 60 479421 ** 6.40 Environment (E) 9 38199744 ** 81.06 GEI 135 494070 ** 15.73 IPCA1 23 1144351 ** 39.46 IPCA2 21 552136 ** 17.38 Residual 91 316313 ns Error 318(300) 281383 246457 * &**= significant at 0.05 and 0.01 probability levels, ns=Non significant, GEI=Genotype by environment interaction, GL=Genotype by location interaction, GY=Genotype by year interaction, LY=Location by year interaction, GLY=Genotype by location by year interaction, (300)=Degree of freedom for error term of AMMI model, SS=Sum of squares. Stability Analysis AMMI Biplots analysis AMMI biplots are recently preferred biplots to visualize adaptability and stability of genotypes over test environments (Gauch and Zobel, 1988; Gauch and Zobel, 1996; Gauch, 2006; Gauch et al., 2008). This is because they are effective tools to diagnose the GEI patterns graphically. In AMMI1 biplot, the genotypes with IPCA1 scores close to zero express general adaptation and the larger scores depict more specific adaptation in combination with environments of the same sign IPCA1 scores (Ebdon and Gauch, 2002a). Furthermore, the relative magnitude and direction of genotypes along the abscissa and ordinate axis in biplot is also important to understand the response pattern of genotypes across environments and to
  • 5. Seed Yield Stability and Genotype x Environment Interaction of Common Bean (Phaseolus vulgaris L.) Lines in Ethiopia Int. J. Plant Breeding Crop Sci. 138 Figure 1. AMMI 1 biplot showing mean performance and adaptability of small red common bean lines over environments differentiate high yielding and adaptable genotypes (Samonte et al., 2005). Accordingly, in Figure 1, G2, G1, and G5 placed relatively close to zero IPCA1 score line and performed above the overall mean were generally adapted to all small red common bean lines growing environments of Ethiopia. The high yielding line (G8) had similar sign IPCA1 scores with IPCA1 score sign of HRM13 and ALT13 showed positive interaction with these environments. Thus, it was specifically adapted to these environments and similar agro-ecologies. Similarly, the high yielding lines (G15 and G9) with similar sign of IPCA1 score to MLK13, MLK14, and HRM14 showed positive interaction with these environments and specifically adapted to these and similar small red common bean cultivars growing environments of Ethiopia. Considering environments, MLK14, MLK13 and HRM14 exhibited high seed yield performance (Fig. 1). Thus, they are better environments for commercial production of common bean lines found specifically or widely adapted to them. MIS13, and MIS14 were low seed yield potential environments. As it is located furthest away from zero line of IPCA1 score, MLK14 showed greatest interaction with lines. This could be due to the less fertile soils of Miesso and better moisture of Melkassa and Haramaya (Table 1). In overall, the lines adaptability/stability ranking for seed yield performance based on lower absolute IPCA1 score value which measures the distance of each line from zero value line, i.e., absolute adaptability line, was G13 (0.33) > G4(1.04) > G5 (2.37) > G2 (3.01) > G3(4.27) > G11 (4.67) > G1(6.16) > G14 (7.46) > G12 (9.17) > G10 (9.44) > G8 (11.28) > G9(11.56 ) > G7(136.91) >G15 (17.86) > G16 (39.29) (Fig. 1). This indicated that G13 was the most stable small red common bean line , but below average seed yielding and adapted to low seed yielding location Miesso and similar agro-ecologies while G16 was the most unstable, low seed yielding, and un- adapted to Ethiopa compared to the lines in the test set. The AMMI1 biplot, Fig. 1, had visualized not only the lines performance in relation to adaptability and mean seed yield performance, but also revealed presence of two mega-environments. Five low and medium seed yield potential environments (MIS13, MIS14, ARN13, HRM13, and ALT13) with similar IPCA1 scores had formed one mega-environment whereas another five medium and high seed yield potential environments (ALT14, ARN14, MLK14, HRM14, and MLK13) had formed another mega- environment. However, its mega-environment classification is more general and didn’t show detailed specific adaptation of lines (Yan et al., 2007). Therefore, more specific adaptability of lines was explored using AMMI 2 biplot (Fig. 2). In AMMI 2 biplot, Figure 2, the small red common bean lines (G16, G5, G13, G15, and G7) placed furthest away from the biplot origin expressed a highly interactive behavior (positively or negatively) whereas G3 and G4 placed relatively close to the biplot origin expressed less interaction and more adapted to all environments. This is because, in AMMI 2 biplot, the distances from the biplot origin are indicative of the amount of interaction exhibited by genotypes over environments or environments over genotypes and genotypes located near the biplot origin are less responsive than the vertex genotypes indicating general adaptability to all environments (Voltas et al.,2002). Environments, MLK 14 and MIS 14, with longer vectors were very interactive and discriminated the differences among genotypes more than other environments with shorter vectors, which were less interactive and provided little information about the
  • 6. Seed Yield Stability and Genotype x Environment Interaction of Common Bean (Phaseolus vulgaris L.) Lines in Ethiopia Ashango et al. 139 Figure 2. AMMI 2 biplot showing general and specific adaptability of small red common bean lines over environments Figure 3. Genotype focusing scaled GGE scatter biplot showing mean performance and stability of small red common bean lines. differences among the lines' seed yield performances as reported by (Yan, 2002). Similarly, in Figure 2, the angles between genotype, environment, or between genotype and environment vectors determine the nature of GEI. The interaction is positive for acute angles, zero for right angles, and negative for obtuse angles (Kandus et al., 2010). Accordingly, G8, G11, and G5 which made acute angles with ARN13 and ALT13 vector showed positive interaction and specifically adapted to these environments. The lines, G15, G1, G2, G5, made acute angles with ALT14 and MLK14 vectors and interacted highly positively with them , therefore, were specifically adapted to them. G9, G7, and G10 were specifically adapted to MLK13. Again, G7, G9 and G6 interacted highly positively with ARN14 and HRM14 and specifically adapted to them. The low seed yielding line, G13, had made acute angle with low seed yield potential environments, MIS13 and MIS14, vectors was specifically best line for them. MIS13 being placed close to the biplot origin showed less interaction with lines than other environments and provided little information about lines' seed yield performance. The low seed yielding line, G16, being placed furthest away from all environments was un- adapted to all environments compared to the lines in the test set. GGE Biplots Analysis Mean Seed Yield Performance and Stability of Lines In GGE scatter biplot scaled focusing genotype, PC1score estimates mean yield with its zero line indicating average performance while the least absolute PC2 score shows top stability (Yan and Kang, 2002). Thus, in Figure 3, lines which had PC1 scores greater
  • 7. Seed Yield Stability and Genotype x Environment Interaction of Common Bean (Phaseolus vulgaris L.) Lines in Ethiopia Int. J. Plant Breeding Crop Sci. 140 Figure 4. Average environment coordination (AEC) view of the GGE comparison biplot showing performance of lines compared with best line performance. than zero viz, G15, G1, G9, G6, G10, G7, G2, and G5 were higher seed yielding lines while lines with PC1 scores less than zero, G8, G11, G3, G14, G13, G4, G12, and G14 were lower seed yielding lines. G14 placed closest to zero line of PC2 score was the most stable, but low seed yielding. Lines placed furthest away from zero PC2 line were unstable. In genotype focusing scaled GGE ranking biplot, Figure 5, lines, G16, G14, G4, G13, G7, G10, and G12, which had fallen below the AEC ordinate showed below average seed yield performance whereas lines, G15, G1, G6, G9, G2, G5, G11, G8, and G3, which had fallen above the average environment coordination (AEC) ordinate performed above average. G1, G2, G3, G11, and G5 which had performed above average and had relatively shortest projection vectors from AEC line were both high seed yielding and widely adapted. On the other hand G4, G13, and G14 which had fallen below AEC ordinate axis and with shortest projection vectors from AEC line were widely adapted, but low seed yielding. This is because, in genotype focusing scaled ranking GGE biplot, AEC approximates the genotypes' main effect, that is, mean performance with the arrow pointing to greater genotype main effect while its ordinate approximates GEI effects, that is, stability with increasing GEI effects and instability away from the origin at both directions (Yan et al., 2001; Yan and Hunt, 2002). In genotype focusing scaled GGE comparison biplot, Figure 4, line, G1, is ideal line and G5, G2, G15, G11, and G8 placed relatively closest to G1 were both higher seed yielding and widely adapted lines compared to other lines. In genotype focusing scaled comparison GGE biplot, a genotype located nearest to the central concentric circle is both high seed yielding and most stable. It is considered as an ideal genotype and genotypes fall closest to it are also considered as desirable (Yan, 2002). In symmetrically scaled polygonal GGE biplot, Figure 6, seven sectors of which four with environments were observed. MLK14, ALT14, HRM14, ARN14, and MIS13 were clustered in one sector and could be considered as one mega-environment for small red common bean lines evaluation and recommendation. Their higher seed yielding lines were G15, G1, G5, and G2. ALT13 and ARN13 had entered into one sectors had also formed another mega-environment with their higher seed yielding lines G8, and G11. MLK13 alone made one mega- environment and its higher seed yielding lines were G7 and G10. MIS13 and HRN13 had made fourth mega- environment and their winning lines were G12 and G3. These specific adaptability and mega-environments interpretations are because, in symmetrically scaled polygon view of GGE biplot, connecting the extreme genotypes forms a polygon and the perpendicular lines to the sides of the polygon form sectors of genotypes and environments, which are considered as separate mega- environments (Gauch and Zobel, 1997; Yan et al., 2000). GEI reflects differences in adaptation and can be exploited by selecting for specific adaptation if the trend in specific adaptability of genotypes is repeatable over years (Annicchiarico, 2002; Yan et al., 2007). However, in this study, the specific adaptability trend was not repeated over years as different environments were grouped differently in two years (Figures 7). Therefore, GEI couldn't be exploited and should be minimized by selecting for broad adaptation. Thus, broadly adapted lines, G1, G2, and G5, were recommended for verification and release.
  • 8. Seed Yield Stability and Genotype x Environment Interaction of Common Bean (Phaseolus vulgaris L.) Lines in Ethiopia Ashango et al. 141 Figure 5. Average environment coordination (AEC) view of the GGE ranking biplot showing mean performance and stability of lines. Discriminating ability, Representativeness, and Relationships of test Environments In environment focusing scaled vector view of GGE biplot, the cosine of the angles between environment vectors show relationships between test environments with acute angles indicating strong correlation, obtuse angles strong negative correlation or cross over GEI of genotypes, and right angle showing no correlation (Yan and Tinker, 2006). Hence, in Figure 8 right, (ALT13 and ARN13) and (MLK14 and ALT14) with acute angles between them were strongly correlated and indicated significant influence of years on genotypes' seed yield performance. Thus, similar information could be obtained by dropping either of one environment for small red common bean lines evaluation by reducing cost of multi- location replicating trials. Therefore, it is better to drop Alem Tena from trial sites as similar information can be obtained from Melkassa and Arsinegelle. MLK14 with the longest vector length was the most discriminating and more Figure 6. Symmetrically scaled polygonal GGE biplot showing specific adaptability of lines.
  • 9. Seed Yield Stability and Genotype x Environment Interaction of Common Bean (Phaseolus vulgaris L.) Lines in Ethiopia Int. J. Plant Breeding Crop Sci. 142 Figure 7. Symmetrically scaled polygonal GGE biplots showing specific adaptability of lines during 2013 (right) and 2014 (left). Figures 8. Environment focusing scaled vector view of GGE biplot (right) and average environment coordination (AEC) view of GGE biplot (left) showing relationships among test environments. informative environment while MIS13 with the shortest vector was the least discriminating and less informative environment. The lines interaction with environments was crossover in MLK14 and MIS14 as indicated by obtuse angle between their vectors (Fig 8, right). Similarly, in environment focusing scaled comparison GGE biplot (Fig. 8 left), a test environment with smallest angle between the AEC abscissa is the most representative (Yan and Tinker, 2006). Hence, MLK14 laid on the AEC abscissa line was the most
  • 10. Seed Yield Stability and Genotype x Environment Interaction of Common Bean (Phaseolus vulgaris L.) Lines in Ethiopia Ashango et al. 143 representative of all environments followed by ALT14. Therefore, MLK14 was both most representative and most discriminating. This is because Melkassa is nearly optimum environment in terms of all edaphic, climatic, and biotic seed yield limiting factors, but Miesso is characterized by less fertile soils, low moisture availability, and higher temperature. Alem Tena, Arsinegelle, and Haramaya are average on all seed yield limiting factors and are medium seed yielding environments. CONCLUSION GEI is differential phenotypic performance of genetically uniform genotypes across test environments. It occurs because different genotypes have varying genetic potentials to adjust themselves to variable environments. Small red common bean lines evaluated by this study had highly significant genetic differences for seed yield performance across environments. Spatial variation in environments was more profound than temporal variations in exerting effects on lines' seed yield performance. AMMI 1, GGE ranking, and GGE comparison biplots enabled identification of broadly adapted and high seed yielding lines better than AMMI 2 and polygonal GGE biplots. Small red common bean lines, KG-71-1, KG-71-23, and KG-71-44 were both high seed yielding and broadly adapted to dry bean growing environments of Ethiopia. AMMI 2 and polygonal GGE biplots enabled selection of specifically adapted lines, but the specific adaptability show of polygonal GGE biplot is clearer than AMMI 1 and AMMI 2 biplots. Environment focusing scaled GGE biplots also enabled identification of ideal (Melkassa), least informative (Miesso), and redundant (Alem Tena) environments for small red common bean lines evaluation. 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