Study on Genotype X Environment Interaction of Seed Yield, Oil Content, Fatty Acid Profile and Stability Analysis of Yield Related Trait in Linseed (Linum usitatissimum L.) in North Western Ethiopia
Linseed (Linum usitatissimum), or common flax, is a useful plant that has been cultivated for thousands of years. Its oil is rich in omega-3 fatty acids, especially Alpha-Linolenic Acid (ALA) (C18:3) that was beneficial for heart disease, inflammatory bowel disease, arthritis and a variety of other health conditions. Nine linseed (Linum usitatissimum L.) genotypes were evaluated in randomized complete block design with three replications during 2011 main cropping season at six locations to determine the pattern of genotype by environment interaction of yield and yield related traits to identify the most stable linseed genotypes for wide and/or narrow adaptations. Combined analysis of variance (ANOVA) indicated significant variations among environments and among genotypes, indicating the existence of variability among the tested genotypes. Debark was the most suitable environment for seed yield, oil content, oil yield and most important fatty acid alpha-linolenic acid. Genotype by Environment Interaction (GEI) was statistically significant only for days to flowering, days to maturity and seed per capsule, indicating the importance of stability analysis. Additive main effect and multiplicative interaction (AMMI) stability analysis in this study selected Kulumsa-1 as stable genotypes for seed per capsule. Linseed adapts to North West Amhara Region, Ethiopia although differences were observed among genotypes. According to the combined analysis of variance for seed yield, oil content, oil yield and alpha-linolineic acid variety Kulumsa-1 can be used for wider cultivation to the North West Amhara Region, Ethiopia.
to cite this article:
Cherinet Alem and Tadesse Dessalegn, 2014. Study on Genotype X Environment Interaction of Seed Yield, Oil Content, Fatty Acid Profile and Stability Analysis of Yield Related Trait in Linseed (Linum usitatissimum L.) in North Western Ethiopia. International Journal of Plant Breeding and Genetics, 8: 66-73.
Received: December 24, 2013;
Accepted: January 28, 2014;
Published: April 15, 2014
Linseed (Linum usitatissimum), or common flax, is a useful plant that
has been cultivated for thousands of years. It is grown for both of its fiber
and its seeds. The fiber from its stems is used to make linen cloth, while linseed
oil is derived from its seeds. Flax seeds are also edible and contain important
nutrients. Linseed oil is rich in omega-3 fatty acids, especially alpha-linolenic
acid (C18:3) that was beneficial for heart disease, inflammatory bowel disease,
arthritis and a variety of other health conditions. It also contains a group
of chemicals called lignans that play a significant role in the prevention of
cancer. Ethiopia is considered to be the secondary center of diversity, and
now the 5th major producer of linseed in the world after Canada, China, United
States and India (Wakjira, 2007). Linseed has long history
of cultivation by smallholders, exclusively for its oil in the traditional agriculture
of Ethiopia (Belayneh and Nigussie, 1988). It is usually
cultivated in higher elevations where frost is a threat for other oil seed crops.
It is a major oil seed and second most important oil crop after noug seed (Guizotia
abyssinica Cass.) in Ethiopia. The crop performs best in altitudes ranging
from 2200-2800 masl; but it is also found in areas as low as 1200 masl and as
high as 3420 masl. It is an important rotational crop for cereals and pulses
Linseed requires cool temperature during its growing period for better yield.
It grows well within temperature ranges of 10-30°C but it performs best
between 21-22°C. It prefers dry and sunny weather with well-distributed
moderate rain over the growing season (Getinet and Nigussie,
1997). Optimum soils for flax are well drained but moisture retentive and
medium to heavy textured, such as clay loams and silty clays. The soil should
be of a fine tilth and not prone to crusting. Flax will not perform well on
soils with pH less than 5 or above 7 and is sensitive to soil salinity (Wakjira,
Worldwide, linseed total area harvested during the year 2010 was 2218625 ha;
with production of 1922759 tons, average yield of 866.6 kg ha-1 and
linseed oil production of 613944 tons (FAO, 2012). In
Africa, linseed total area harvested during the year 2010 was 153431 ha; with
seed production of 166690 tons, average yield of 1086.4 kg ha-1 and
linseed oil production of 52009 tons (FAO, 2012). In
Ethiopia, its total area during the year 2008 was 180872.7 ha; with the total
production of 156079 tons and average yield of 0.863 tons ha-1 (CSA,
2008). In Ethiopia linseed is produced mostly for edible oil. But the remains
after oil extraction are used for animal feed (Getinet and
Nigussie, 1997). So far several varieties of linseed have been released
in Ethiopia by national and regional research institutions (MoARD,
It is commonly observed that the relative performance of different genotypes
varies in different environments, i.e., there exists genotype-environment interaction.
Presence of significant genotype by environment interaction due to the differential
response of varieties in different environments represents a major challenge
to plant breeders to fully understand and obtain the genetic control of variability
(Luthra and Singh, 1974). In this case, measuring and
understanding the Genotype by Environment Interaction (GEI) should be an essential
component of variety evaluation. One of the main reasons of growing varieties
in multilocations is to estimate their stability (Freeman,
1973) as selection of superior varieties is mainly based on their yield
potential and stable performance over a wide range of environments (Crossa
et al., 1989).
An understanding of the genetic and environmental basis of genotype by environment
interaction is of fundamental importance in plant breeding (Crossa
et al., 1999). Mean seed yield is an important criterion for the
selection of superior varieties in the absence of GEI. However, a stability
analysis is necessary when GEI is significant and its contribution to the total
sums of squares is higher than that of genotypes contribution. Successful linseed
cultivars must perform reliably in yield and other agronomic traits over wide
range of environments (Diepenbrock et al., 1995).
In Ethiopia, there are only few studies reported on the genotype by environment
interaction and stability of linseed varieties in the past. One study by Adugna
and Labuschagne (2002) indicated that year by location and location variability
was found to be dominant sources of interactions for linseed in Ethiopia. But
further investigation on GEI and stability of linseed in Ethiopia is crucial
to accumulate more scientific evidences for any anticipated changes like climatic
irregularities. To date, little information is available on this crop and its
adaptation pattern, especially under Northwestern Ethiopian conditions. Keeping
this in view, the present study was conducted to examine the pattern of GEI
of yield and yield related traits; to identify the most stable linseed variety
for wide and/or specific adaptations.
MATERIALS AND METHODS
Nine released linseed genotypes (CI-1525, CI-1652, Chilallo, Belay-96, Geregera,
Berene, Tole, Kulumsa-1 and Geldu) were evaluated at six different locations
during 2011 main cropping season.
|| Brief description of experimental sites
|Ethiopian Metrological Agency, Bahir Dar branch, NA: Data
The detailed information about locations is presented in Table
1. Randomized Complete Block Design (RCBD) with three replications was used
throughout the testing locations. Each experimental plot had six rows of 5 m
l ength and 20 cm spacing between rows. A seed rate of 45 kg ha-1
was used by hand drilling the seeds in the rows. Fertilizers rates of 50/30
kg ha-1 DAP and Urea respectively was used for all sites and applied
fully at time of planting. Planting was carried out from mid to the end of June
2011 following the farmers' practice. All other recommended agronomic and cultural
practices were carried out for all the plots uniformly. Twenty different data
were collected from the middle four rows. Combined analyses of variance over
locations were done using AGROBASE 20 software 2000.
RESULTS AND DISCUSSION
Combined analysis of variance (not presented) showed highly significant variations
among environments for all characters considered except plant height and among
genotypes across all locations for days to flower, days to maturity, plant height,
thousand seed weight, oil content, palmitic acid, stearic acid, oleic acid,
linoleic acid, linolenic acid, tiller per plant and capsule per plant, indicating
the existence of variability among the tested genotypes. Significant variations
among locations for days to flower, days to maturity, seed per capsule, branch
per plant and capsule per plant and among genotypes for days to flower, days
to maturity, plant height and thousand seed weight were also reported by Adugna
and Labuschagne (2003). Significant variation among genotypes for oil content
was also reported by Wakjira et al. (2004).
Ahmed and Ahmed (2012) in sesame; Patil
et al. (1999) in safflower and Choferie (2008)
in linseed reported significant differences between genotypes for days to flowering
and days to maturity. Berti et al. (2010) also
reported significant differences between genotypes for oil content and stearic,
oleic and linolenic acid in linseed. Similarly, Gabiana (2005)
reported significant differences between genotypes for oleic, linoleic and linolenic
acid in linseed. Significant genotype by environment interaction was observed
only for days to flower, days to maturity and seed per capsule. Similarly, significant
GEI for days to flower and days to maturity was also reported by Adugna
and Labuschagne (2003) and Choferie (2008). It agrees
with the finding that yield and agronomic traits are influenced by genotypes,
environment factors and the interaction between genotype and environment (Adugna
and Labuschagne, 2003; Wakjira et al., 2004;
Choferie, 2008; Berti et al.,
2010; Gunasekera et al., 2006; Mostafa
and Ashmawy, 1998).
Analysis of variance (ANOVA): The analysis of variance of seeds per
capsule of nine genotypes tested in six environments is presented in Table
2 and 3. The analysis revealed that linseed genotypes
were significantly (p<0.01) affected by Environments (E) and Genotype by
Environment Interaction (GEI).
||Additive main effects and multiplicative interaction analysis
of variance for seeds per capsule of the genotypes across environments
|*, **Significant at 5 and 1%, respectively, Env: Environments,
DF: Days to flower, SS: Sum of square, MS: Mean sum of square
||Mean seed yield (kg ha-1), Thousand Seed Weight
(TSW), oil content (%), oil yield (kg ha-1), seed per capsule
and fatty acid profiles of 9 linseed genotypes tested at 6 locations in
the GEI study (2011-12)
|*, **Significant at 5 and 1%, respectively. SY: Seed yield
(kg ha-1), TSW =1000- Seed weight (g), OC: Oil content (%), OYH:
Oil yield (kg ha-1), SPC: Seeds per capsules and fatty acid profiles
The main effects of Environments (E) and Genotype by Environment Interactions
(GEI) accounted for 52.97 and 13.42% of the total variation in GxE data for
seeds per capsule. High variability among environments and genotypes existed
both in the main and interaction effects for seeds per capsule. Potential environments
for seed per capsule were distributed in the 4th quadrant (Lai-Gaint), and in
the 1st quadrant (Adet, Debark, Debretabor and Merawi), whereas the low potential
environment Dabat (B), was distributed in the 2nd quadrant (Fig.
1). Environments A (Adet), C (Debark), D (Debretabor), E (Lai-Gaint) and
F (Merawi) gave above average mean seed per capsule, while location B (Dabat)
gave below average (Fig. 1). Locations A (Adet), C (Debark),
D (Debretabor) and F (Merawi) were in the same quadrant (I) for seed per capsule
and hence they discriminated between the genotypes in much similar ways, while
location E (Lai-Gaint) differed from these high potential environments hence
discriminated between the genotypes in different way.
||AMMI biplot of mean seed per capsule and the first Principal
Component Axis (PCA) 1 scores of 9 linseed genotypes at six environments
in 2011, Genotypes plotted as a, b, c ... and environments as A, B, C...,
a: CI-1525, b: CI-1652, c: Chilallo, d: Belay-96, e: Geregera, f: Berene,
g: Tole, h: Kulumsa-1, i: Geldu, A: Adet, B: Dabat, C: Debark, D: Debretabor,
E: Lai-Gaint, F: Merawi
|| Environment means of seed per capsule and their EIPC1 and
EIPC2 scores of six locations for the 9 linseed genotypes
Location E (Lai-Gaint) had the highest interaction with the genotypes, followed
by F (Merawi) and A (Adet) because they had higher EIPC1 scores for seed per
capsule (Table 4).
Genotypes CI-1652 (b), Belay-96 (d), Tole (g) and Geldu (I) had positive interaction
with the potential environment, Lai-Gaint (E), while these genotypes had negative
interaction with the potential environments, Adet (A), Debark (C), Debretabor
(D), Merawi (F) and with the low potential environment, Dabat (B) for seed per
capsule. Genotypes CI-1525 (a), Chilao (c), Geregera (e), Berene (f) and kulumsa-1
(h) had positive interaction with the potential environments, Adet (A), Debark
(C), Debretabor (D), Merawi (F) and with the low potential environment, Dabat
(B), while these genotypes had negative interaction with the potential environment,
Lai-Gaint (E), for seed per capsule (Table 5).
Varieties Geldu, CI-1652 and kulumsa-1 had absolute IPCA1 scores very close
to zero for seed per capsule. CI-1652 and kulumsa-1 had also mean seed per capsule
above average and these were the most stable varieties across location (Table
When biplot interaction for the AMMI2 model (Fig. 2) is considered,
genotypes Geldu, CI-1652, Kulumsa-1 and CI-1525 were the most stable varieties
for seed per capsule; while Geregera and Berene were moderately stable for seed
per capsule. On the other hand, genotypes Chilalo, Belay-96 and Tole were found
unstable for seed per capsule.
||Biplot of Principal Component Analysis axis (PCA) 1 against
principal component analysis axis (PCA) 2 for seed per capsule of 9 linseed
genotypes at six environments in 2011, Genotypes plotted as a, b, c ...
and environments as A, B, C... a: CI-1525, b: CI-1652, c: Chilallo, d: Belay-96,
e: Geregera, f: Berene, g: Tole, h: Kulumsa-1, i: Geldu, A: Adet, B: Dabat,
C: Debark, D: Debretabor, E: Lai-Gaint, F: Merawi
||Scores of genotypes and locations to the first interaction
principal component axis for days to flowering, days to maturity and seed
|SPC: Seed per capsule, DF: Days to flower, DM: Days to maturity
Generally, the present study entails the presence of significant variations
among environments and among genotypes for most of the characters, indicating
the existence of variability among the tested genotypes. Linseed adapts to Northwest
Amhara region, Ethiopia, although differences were observed among genotypes
for days to flowering, days to maturity, plant height, thousand seed weight,
capsule per plant, and oil content and composition. Seed yield was above 1700
kg ha-1 for some environments and genotypes, indicating the high
seed yield potential of some environments in Northwest Amhara region, Ethiopia.
Debark was found to be the most suitable environment for seed yield, oil content,
oil yield and most important fatty acid alpha-linolenic acid. According to the
combined analysis of variance for seed yield, oil content, oil yield and alpha-linolineic
acid variety Kulumsa-1 can be used for wider cultivation to the to Northwest
Amhara region, Ethiopia.
Special thanks goes to Adet Agricultural Research Center (AARC) and Public
Private Partnership Project from Netherland Embassy (PPPO) for the financial
assistance and resources. I am also highly thankful to Bahir Dar University
for educating me to the level of M.Sc. degree.
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